TELOPEA
Volume 8(4) • July 2000
Royal Botanic Gardens Sydney
National Herbarium of New South Wales
Telopea is published by the National Herbarium of New South Wales, Royal Botanic
Gardens Sydney. It covers the field of systematic botany in Australia specialising in the
flora of New South Wales.
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Cover illustration
The Waratah, Telopea speciosissima (Sm.) R. Br., belongs to the family Proteaceae. The
species is endemic in eastern New South Wales and is the official State floral emblem.
Illustration by David Mackay
Telopea
A journal of plant systematics
CONTENTS
A new species of Minuria (Asteraceae: Astereae) from
New South Wales P.S. Short and J.R. Hosking 407
A revision of Davidsonia (Cunoniaceae)
Gwen J. Harden and John B. Williams 413
Seedling characteristics in the Casuarinaceae
Robin Hiuang and John G. Conran 429
The taxonomy of Xanthosia liuegelii and closely related species
(Apiaceae: Hydrocotyloideae) /. M. Hart 441
Actinotns periculosus (Apiaceae): a new perennial species from
eastern Australia M.J. Henwood 455
The lichen genera Phaeographis and Phaeographina (Graphidaceae)
in Australia 1: Species based on Australian type specimens
Alan W. Archer 461
Hopkinsiaceae and Lyginiaceae, two new families of Poales
in Western Australia, with revisions of Hopkinsia and Lyginia
Barbara G. Briggs and L.A.S. Johnson 477
Systematic studies in the eucalypts. 10. New tropical and subtropical
eucalypts from Australia and New Guinea ( Eucalyptus, Myrtaceae)
K.D. Hill and L.A.S. Johnson 503
TELOPEA 8(4): 407-539 JULY 2000
Scientific Editor
Peter Wilson
Editor of Publications
Gary Bridle
Typesetting and Production
Debby McGerty
Other members of Editorial Committee
Tim Entwisle, Joy Everett, Alistair Hay
ISSN 0312-9764
Telopea 8(3) was distributed on 16 December 1999.
National Herbarium of New South Wales
Royal Botanic Gardens Sydney
Mrs Macquaries Rd
Sydney NSW Australia 2000
407
A new species of Minuria (Asteraceae:
Astereae) from New South Wales
P.S. Short and J.R. Hosking
Abstract
Short, P.SJ and Hosking, J.R. 2 PNorthern Territory Herbarium, Parks & Wildlife Commission of the
Northern Territory, P.O. Box 496, Palmerston, NT 0831, Australia; 2 NSW Agriculture, RMB 944,
Tamwortli, NSW 2340, Australia). A new species o/Minuria (Asteraceae: Astereae)from Nezo South Wales.
Telopea 8(4): 407-411. Minuria scoparia, a species restricted to serpentinite-derived soils from
northern New South Wales, is described and illustrated. It has affinities with M. cunninghamii and
M. leptophylla but is broom-like in habit and the pappus bristles of the disc florets are not
manifestly dimorphic.
Introduction
At least six species of plant are apparently endemic to the serpentinite-derived soils
which occur between areas east of Bingara and east of Chaffey Dam, northern New
South Wales (Hosking & James 1998). One of these, a species of Minuria, is
described herein.
The circumscription of Minuria is problematic and unpublished results of cladistic
analyses (by PSS) of the Australasian Astereae indicate that, as currently delimited
(Lander & Barry 1980), the genus is not monophyletic. The species described here
appears to belong to Minuria s. str., its affinities seemingly being with M. leptophylla
DC., the type species of the genus, and M. cunninghamii (DC.) Benth.
Taxonomy
Minuria scoparia P.S.Short & J.R.Hosking, sp. nov.
M. cunninghamii et M. leptophyllae similis sed ab eis habitu scopulato et flosculis discis
pappum manifeste dimorphum deficiens differt.
Holotype: slopes of Blackfellows Knob, east of Chaffey Dam, 31°23'14"S, 151°09'02"E,
Altitude 670 m, J.R. Hosking 1532, 23 Nov 1997 (DNA 134923). Isotypes: CANB, MEL,
NE, NSW.
Perennial, stiff, broom-like, sprawling to erect herb with branches to c. 60 cm long;
major axes mainly glabrous but with whitish, somewhat curled and apically tapering,
eglandular, uniseriate, septate hairs present, especially immediately below the
capitula, a few short, glandular hairs also present. Leaves alternate, sessile although
somewhat constricted and paler at base, entire, linear, (1.5-)12-28 mm long,
(0.3-)0.5-1.2 mm wide, glabrous or with scattered glandular and eglandular hairs,
mucronate. Capitula solitary, heterogamous, radiate. Involucre 4.5-6 mm diam.,
multiseriate. Bracts c. 30-40, lanceolate, 1-3.2 mm long, 0.2-0.4 mm wide, with the
herbaceous stereome divided, the upper margins and apex of the bracts usually
hyaline, mostly green but apex purple. Receptacle flat, glabrous, pitted. Ray florets
female, 29-37, uniseriate, corolla 4.4-5 mm long, the ray 3-veined and apically
408
Telopea 8(4): 2000
minutely bilobed, almost white to pale pink or purple; style branches pink. Cypselas
laterally flattened, symmetrical or almost so, apically barely to manifestly constricted
and sometimes forming a short beak to c. 0.3 mm long; fruit body oblanceolate in
outline, 2-2.4 mm long, 0.5-0.8 mm wide and more or less uniformly brown except for
the carpopodium, with scattered hairs mainly on the margins and at and near the
apex, the hairs eglandular, rigid, biseriate, minutely and slightly unevenly bilobed;
carpopodium distinct, annular, white, of more or less quadrate cells and 4-6 cells
wide; pedicel usually evident emerging from carpopodium. Pappus commonly
appearing monomorphic, consisting of 21-31 uniseriate bristles of about equal length,
the bristles 2-3.1 mm long and uniformly barbellate along their length, united at the
base, sometimes with several minute, few-celled outgrowths at the bases of the main
bristles. Disc florets male, 23-27; corolla actinomorphic, tubular, the tube 2-3.1 mm
long, 5- or rarely 4-lobed, mainly yellow throughout but at least when dry the lobes
pinkish. Anthers 5, 0.9-1.1 mm long, microsporangia 0.7-0.9 mm long, terminal
appendage 0.2-0.25 mm long; pollen grains c. 1500 per floret; style barely divided,
arms more or less oblong, dorsal surface pubescent below the point of bifurcation, the
hairs apically obtuse. Cypselas non-functional, glabrous or with scattered hairs as in
ray cypselas. Pappus generally appearing monomorphic, consisting of 9-22 uniseriate
bristles uniformly barbellate along their length and c. the length of the corolla tube and
united at their bases, sometimes minute few-celled outgrowths present between or
perhaps at the bases of the main bristles. (Fig. 1).
Etymology: the specific epithet reflects the distinctive broom-like habit.
Distribution: confined to the North Western Slopes botanical region of New South
Wales. It is known from four populations, i.e. from the type locality east of Chaffey
Dam, from the site of an old magnesite mine north east of Attunga, from near
Woodsreef asbestos mine and from near Upper Bingara (Fig. 2).
Habitat: apparently restricted to serpentinite-derived, skeletal soils. The type
collection is from an Angophora floribunda-Euccilyptus sp. aff. macrorhyncha woodland
vvhere Acacia paradoxa is the dominant shrub and Triodia scariosa the dominant grass.
1 he Attunga and Upper Bingara localities are similar. The Woodsreef locality is also in
Eucalyptus sp. aff. macrorhyncha woodland and plants grow with grasses and herbs
such as Aristida ramosa var. ramosa, Themeda trumdra, Lepidosperma lalerale, Senecio lautus
subsp. dissectifolius and Cheilanthes distorts. More details of the vegetation of this region
are outlined by Hosking and James (1998).
Conservation status: the populations of M. scoparia fall within a stretch of
approximately 160 km. The population east of Chaffey Dam is on government land
vested in the Department of Land and Water Conservation, that at Woodsreef is on
Crown Land, and those near Attunga and Upper Bingara are privately owned. All four
are in areas that, due to steep slopes, are not often grazed by cattle or sheep. At the
type locality and at Attunga and Upper Bingara the presence of Triodia scariosa also
fnn erS t 8 w Z1 j 8 ' M f re than 50 plantS Were observed at th e type locality and more than
I°ll!! 0 . 0dSree ' Tens ° f thousands were observed at the locality near Attunga
g° 52 02 S ' | 50 ° 54 37 " E ' altitude 550 m). Only four plants were observed at the Upper
Bingara locality (30°01'23"S, 150°37T7"E, altitude 590 m). The species is also likely to
be found at other serpentinite localities between Bingara and Chaffey Dam.
Although it is restricted in distribution the species is in no way endangered and given
the size of the population near Attunga it should not be considered rare.
Flowering period and breeding system: the species appears to flower for most of
year, flowering being recorded for April, June, August and November.
the
Short and Hosking, A new species of Minuria (Asteraceae: Astereae)
409
Fig. 1. Minuria scoparia. a, habit; b, mature receptacle; c, disc floret; d, mature cypsela showing
prominent beak and pappus; e, apex of mature cypsela with short beak; f, apex of mature cypsela
with unformed beak, (a, b from Hosking 1532; c-f, from Hosking 1578). Scale bars: a = 3 cm;
b-f = 2 mm.
410
Telopea 8(4): 2000
Fig. 2. Distribution of Minuria scoparia.
A pollen:ovule ratio of 849 was determined from a single capitulum containing 37
female and 24 male florets. This value and the relatively inconspicuous ray florets
(compared to other Astereae with ray florets) suggests that the species commonly self-
pollinates and is self-compatible (e.g. Short 1981, Watanabe et al. 1991).
Notes: apart from recent collections (by JRH in 1992, 1997 & 1998), the only other
specimens known to us were gathered by the Reverend H.M.R. Rupp from Woodsreef
in April and August 1913. His collections are housed in NSW. Tire most distinctive
feature of the species, its habit, is not evident from Rupp's specimens and they were
at one stage incorporated under the name Vittadinia australis A.Rich. var. tenuissima
Benth. and subsequently under the name Minuria cunninghamii. It is with the latter
species and M. leptophylla that M. scoparia is most likely to be confused. It is readily
differentiated from both, not only by its broom-like habit but also by differences in
pappus morphology. In M. scoparia the pappus in both ray and disc florets generally
appears to be monomorphic, with occasionally in both types of floret few-celled
outgrowths occurring at the base of the large bristles. In both M. cunninghamii and
M. leptoplnjlla the pappus of the disc florets is manifestly dimorphic as illustrated in
Lander (1992).
Short and Hosking, A new species of Minuria (Asteraceae: Astereae)
411
The above description is based on the holotype specimen from Blackfellows Knob,
duplicates of Hosking 1578 from Woodsreef that are held at AD and DNA, and notes
supplied by JRH to the senior author. The limited sample suggests that both
populations can be differentiated from each other by several morphological
characteristics. At the Woodsreef populations the ray cypselas may be shortly beaked,
the pappus bristles of the ray florets are c. 3 mm long, there are 14-22 pappus bristles
per disc floret and the corolla tube of the disc florets is c. 3 mm long. In the holotype
specimen manifestly beaked cypselas are absent, the pappus bristles of the ray florets
are 2-2.2 mm long, there may be fewer pappus bristles in the disc floret and the corolla
tube in disc florets is c. 2-2.4 mm long. The colour of the ray corollas also differs
between populations. At the type locality the rays are almost white or pale pink, at
Woodsreef they are pale purple, drying to dark purple.
Specimens examined (excluding type): New South Wales: North Western Slopes: steep slope
above Ironbark Creek, west of Woodsreef asbestos mine, 30°24'H"S, 150°43'24"E, altitude 500 m,
Hosking 1578 & Watts, 19 June 1998 (AD, BRI, CANB, DNA, MEL, NSW, NE); same location, Hosking
635, 25 Nov 1992 (CBG, MEL, NE, NSW); Woodsreef, Barraba, Rupp s.n., Apr 1913 (NSW 122362);
same location, Rupp s.n,, Aug 1913 (NSW 122361).
Acknowledgments
We thank Emma Short for providing the Latin diagnosis, Beth Chandler for the
botanical illustration and Paul Munns for the map. We also thank James Fensbo for
access to the magnesite mine area northeast of Attunga and various land managers in
the Department of Land and Water Conservation for access to serpentinite areas east
of Chaffey Dam and at Woodsreef.
References
Hosking, J.R. & James, T.A. (1998) Native and exotic flora of the North Western Slopes upstream
of the junction of the Peel and Namoi Rivers, New South Wales. Cunninghamia 5: 721-766.
Lander, N.S. (1992) Minuria. Pp. 167-169 in Harden, G.J. (ed.). Flora of New South Wales, vol. 3.
(New South Wales University Press: Kensington).
Lander, N.S. & Barry, R. (1980) A review of the genus Minuria DC. (Asteraceae, Astereae). Nuytsia
3: 221-237.
Short, P.S. (1981) Pollen-ovule ratios, breeding systems and distribution patterns of some Australian
Gnaphaliinae (Compositae: Inuleae). Muelleria 4: 395-417.
Watanabe, K., Short, P.S., Kosuge, K. & Smith-White, S. (1991) The cytology of Brachyscome Cass.
(Asteraceae: Astereae). 11. Hybridization between B. goniocarpa Sond. & F. Muell. (n = 4) and
B. dichromosomatica C. R. Carter (n = 2). Austral. /. Bot. 39:475-485.
Manuscript received 20 September 1999
Manuscript accepted 16 February 2000
Note added in proof
After this paper had been accepted for publication an additional population (Hosking
1846, CANB, DNA, MEL, NE, NSW) of M. scoparia, also growing on serpentinite soil,
was found alongside Carters Road east of 'Boonara', 31°47'36"S, 151°36'33"E, altitude
580 m. This is in the North Coast botanical region of New South Wales, approximately
60 km southeast of the type locality at Blackfellows Knob.
413
A revision of Davidsonia (Cunoniaceae)
Gwen J. Harden and John B. Williams
Abstract
Harden, G.J. 1 and Williams, J.B. 2 0National Herbarium of New South Wales, Royal Botanic Gardens,
Sydney, NSW 2000, Australia; 2 Botany Division, University of New England, Armidale, NSW 2351,
Australia), A revision of Davidsonia (Cunoniaceae). Telopea 8(4): 413-428. The genus Davidsonia
(Cunoniaceae) is revised on the basis of available herbarium collections, field studies over its
geographic range, fresh and preserved materials and previous accounts. Formerly considered a
monotypic genus, Davidsonia is found to comprise three species; D. pruriens F. Muell., the type
species, from north-eastern Queensland; D. jerseyana (F. Muell. ex F.M. Bailey) G. Harden & J.B.
Williams, a known variety from north-eastern New South Wales here elevated to specific rank; and
D. johnsonii J.B. Williams & G. Harden, a well-known but hitherto undescribed species from north¬
eastern New South Wales and south-east Queensland. Each species is fully described and
illustrated, and the lectotypification of D. pruriens is clarified.
Introduction
When describing tire genus Davidsonia in 1867, Ferdinand von Mueller referred it to
the family Saxifragaceae. Later, when the Cunoniaceae were separated from that
family, Davidsonia was traditionally placed in the Cunoniaceae, though Engler (1930)
urged its exclusion because of the stiff hairs, alternate leaves, short stamens and
exalbuminous seeds. Bange (1952) segregated Davidsonia from the Cunoniaceae into its
own family, the Davidsoniaceae, and this has generally been regarded as a unigeneric
family endemic to eastern Australia. The characters emphasised by Bange (op. cit.)
when he described the family Davidsoniaceae were its spirally arranged leaves, non-
endospermic seeds and epitropous ovules. Cunoniaceae sens, strict, usually has
apotropous ovules but epitropous ovules have also been recorded in Spiraeanthemum
in that family (Dickison 1980). Airy Shaw (1973), Dahlgren (1980), Takhtajan (1980),
Cronquist (1981), and Mabberley (1997) have maintained Davidsoniaceae as a separate
family, closely related to the Cunoniaceae. However, Hutchinson (1973) retained the
genus in the Cunoniaceae without comment, while Ingle and Dadswell (1956) and
Dickinson (1980) argued that the genus should remain in the Cunoniaceae on the
evidence of its wood anatomy. Carpological evidence (Doweld 1998) supports the
segregation of the Davidsoniaceae from the Cunoniaceae; however, the evidence also
emphasises the families' close affinities. Recent research using DNA sequencing
supports the inclusion of Davidsonia in the Cunoniaceae (Bradford pers. comm. 1997,
1999; Angiosperm Phylogeny Group 1998).
The genus Davidsonia was described in 1867 by Mueller with one species, D. pruriens.
The southern populations of D. pruriens were described as a distinct variety, var.
jerseyana, by Bailey in 1900. The brief description of var. jerseyana was that it was
'smaller in all its parts'. The type variety is restricted to north-eastern Queensland, and
the variety jerseyana confined to north-eastern New South Wales, though erroneously
listed by Bange (1952) as also occurring in southern Queensland. These two taxa are
markedly allopatric and differ consistently in inflorescence size and position, anther
shape and size, pyrene morphology and size, arid leaflet venation.
414
Telopea 8(4): 2000
It is considered appropriate that variety jerseyana should be given specific rank. This
follows the evolutionary species concept as outlined by Wiley (1978, 1981), in which
morphologically different allopatric lineages should be given specific rank. A third
taxon occurring in New South Wales and southern Queensland is very distinct from
the above two taxa in several characters and is here described as a new species.
Three collections of Davidsonia (at NSW) made in 1926, 1939 and 1944 from north¬
eastern New South Wales were segregated by L.A.S. Johnson in June 1949 and labelled
'clearly a new species'. We concur with this view and the species is here described as
Davidsonia johnsonii J.B. Williams & G. Harden. D. jolmsonii is largely sympatric with
D. jerseyana (Fig. 4) but no intermediates are known. It differs from the specimens of
D. pruriens var. jerseyana in several characters including growth habit, leaf size, form
and pubescence, and fruit shape and size, and also differs in its reproductive method
in that no seeds are known to be formed. This species was not apparently collected
between 1944 and 1977 when it was rediscovered by Graham Watson growing on his
property at Huonbrook near Mullumbimby, and drawn to the attention of the authors
in 1978. However, a specimen had been collected by A.G. Floyd and FI.C. Hayes in
1958 at The Pocket and deposited at the Coffs Harbour Forestry Herbarium (CFSHB)
and Floyd and Hayes (1961) stated that 'a possible new species at Mooball near
Murwillumbah is quite glabrous'. Since that time collections have been made from
18 localities in north-eastern New South Wales and three in south-eastern Queensland.
The taxon was informally described or listed as a new species in the following
publications (Williams 1979, Williams & Harden 1979, Williams et al. 1984, Floyd
1981,1989 and Harden 1990), with the descriptions of the flowers and fruit in
Floyd (op. cit.) based on information provided by one of the authors (J.B.W.).
In recent years some specific studies at the University of New England have involved
examination of the Davidsonia taxa in New South Wales. They have concentrated on
their conservation and development (Watson 1987) and the pollination of Davidsonia
pruriens var. jerseyana (Elliott 1991). Also, various surveys have been undertaken of
both these taxa in New South Wales to establish the distribution and size of the
populations as part of the ongoing documentation and management of species that are
considered to be rare or threatened (Briggs and Leigh 1988, 1996) or listed in the
schedules of the New South Wales Threatened Species Conservation Act, 1995. These
studies and surveys have either been prepared as thesis requirements for post¬
graduate courses and have not been published, or have been prepared as internal
reports for the National Parks and Wildlife Service of New South Wales (Quinn et al.
1996). One of the authors (J.B.W.) has been involved in each of these studies
and surveys.
Fresh material from plants growing at the Royal Botanic Gardens Sydney was used for
the illustrations. These were supplemented with preserved mature fruits of
D. jerseyana from Mooball.
Davidsonia F. Muell.
Mueller (1867: 4); Bailey (1900: 537); Bange (1952: 294); Harden (1990: 523).
Type species: Davidsonia pruriens F. Muell.
Small trees to 20 m high, usually in rainforest, either with one or a few sparingly
branched stems or a dense-foliaged much-branched canopy; leaves often tufted
towards the end of stems. Indumentum either prominent and persistent on all or most
parts, or inconspicuous or absent on mature parts. Seedling leaves simple, serrate; first
pair of seedling leaves opposite, narrow-elliptic; following seedling leaves alternate.
Harden and Williams, Davidsonia (Cunoniaceae)
415
pinnate with an increasing number of pinnae (3-7), rachis becoming winged.
Adult leaves alternate, imparipinnate, petiolate; rachis winged with the wing or wing¬
like segments dentate; stipules paired, amplexicaul, semi-circular; pinnae sessile or
shortly stalked, narrow-obovate to narrow-ovate in general outline; terminal pinna
usually the largest, with pinnae pairs progressively smaller towards base of leaf;
petiolulate or sessile; stipellae usually present at base of pinnae. Inflorescences
cauliflorous, ramiflorous or subterminal and borne amongst the leaves, paniculate and
open or congested and spike-like. Floivers bisexual, 4- or 5-merous; sepals 4 or 5,
petaloid, pink or red, connate in the basal half, valvate in bud; petals absent; stamens
usually 8 or 10, exserted beyond the sepals, alternating with an equal number of
nectaries; carpels 2 or rarely 3, ovary bilocular or rarely trilocular, ovules 5-7 in each
loculus, styles 2 or rarely 3; ovules anatropous; in one species (D. johnsonii) the flowers
are often sterile, with styles reduced or absent and anthers without pollen. Fruit a
drupe, mesocarp fleshy; pyrenes 2 or rarely 3, compressed, pyrene wall cartilaginous
to semi-woody; body of pyrene with soft fibres extending radially from margins,
dorsally keeled and terminally crested; seed solitary or absent, glossy dark reddish
brown to black, strongly compressed, plano-convex, glabrous, broad-ovate to circular,
apex obtuse with a prominent short point at radicle end; endosperm absent.
The pyrenes lie separately in the fleshy mesocarp and have sometimes been
misinterpreted as seeds (e.g. Bailey 1898, Elliott & Jones 1984).
Three types of trichomes are present: (1) glandular trichomes, multicellular and
multiseriate, capitate, red-tipped, often terminating teeth on bracts, rachis wings,
margins of stipules and pinnae; (2) bristly trichomes (surfaces hirsute, only in
D. pruriens and D. jerseyana) fawn to yellowish or pinkish on young growth, irritant,
often on leaves, stems, rachises, outer surface of sepals and on fruit; (3) short soft hairs
(surfaces softly pubescent) as on the inner surface of sepals (and outer surface in
D. johnsonii).
Key to the species
1 Leaflets, rachis, branchlets, inflorescence axes and outer surface of sepals hirsute, at
least when young, with long irritant bristly hairs (usually moderately to strongly
persistent); most adult leaves more than 35 cm long; pinnae (7—)9—19, the three
largest pinnae (terminal and subterminal) 20-40 cm long; margins of pinnae
secondarily serrate; inflorescences cauliflorous or ramiflorous, usually borne on the
main trunks or sometimes among the leaves; styles 2 or rarely 3, conspicuous,
curved, exceeding the stamens; fruit pyriform to obovoid or ellipsoid, with length
greater than or equal to its diameter.
2 Inflorescences open, paniculate, usually more than 30 cm long; anthers oblong at
anthesis, 1.8-3.0 mm long; pyrenes (when fertile, i.e. containing a seed) 2.0-2.7 cm
long, 2.5-2.9 cm wide with a prominent dorsal crest 3.0-6.0 mm long; adult
leaves usually 60-120 cm long, rachis scarcely winged; upper lateral pinnae
mostly lanceolate to oblong, with 15-27 pairs of secondary veins.
. 1. Davidsonia pruriens
2* Inflorescences condensed, racemose or spike-like, mostly less than 25 cm long;
anthers ellipsoid at anthesis, 1.1-1.7 mm long; pyrenes (when fertile) 2.0-2.2 cm
long, 1.6-1.7 cm wide, with an inconspicuous crest 2.0-2.3 mm long; adult leaves
usually 35-60 cm long, rachis usually prominently winged; upper lateral pinnae
mostly oblanceolate, with 11-17 pairs of secondary veins.
... 2. Davidsonia jerseyana
416
Telopea 8(4): 2000
1* All parts more or less glabrous, sometimes pubescent when young but glabrescent,
hairs not irritant; leaves mostly less than 35 cm long; pinnae 5-9, the three largest
pinnae (terminal and subterminal) usually 10-20 cm long; margins of pinnae
dentate but not secondarily serrate; inflorescences subterminal on branchlets, borne
amongst the leaves; styles reduced or absent, if present 2 or 3, shorter than or
scarcely exceeding the stamens; fruit depressed-globose, with length less than
diameter. . 3. Davidsonia johnsonii
1. Davidsonia pruriens F. Muell.
Mueller (1867: 4, t. XLVI); Bailey (1900: 538); Bange (1952: 294-296).
[Davidsonia pungens was listed in Gardeners' Chronicle in 1876, p.603, but without the
author's name. This was given as a north Australian plant and from the description is
referred to this species.]
Type citation: 'In silvis irriguis ad sinum Rockingham's Bay. Dallachy'
Lectotype: designated by Bange (1952: 296) as Murray's River, Rockingham Bay:
Dallachi/, 12 Aug 1866, a MEL collection with flowers and fruit. As there are nine sheets
at MEL in type folders we propose that MEL 106575 be the lectotype, thus restricting
lectotypification to a single specimen, in line with the recent addition to Article 9 of the
Code. This specimen was annotated by Bange, it consists of portion of a leaf and fruit
and it has a copy of the original notes of Dallachy with the date 12 th August 1866. See
full discussion after species description.
Illustrations: Mueller (1867: t. XLVI), Bailey (1898: t. XXXVII), Bange (1952: 295),
Cronquist (1981: 549).
Small slender rainforest tree to 18 m high; bark brown to dark grey, becoming flaky.
Indumentum prominent on all parts, young growth pinkish, hairs long, irritant; red
gland-tipped trichomes terminating teeth on margins of stipules and stipellae. Seedling
leaves at first simple, margins dentate; becoming more or less lobed at base and pinnate
after 12 months growth; hirsute and with some reddish glandular hairs present on
stipules. Adult leaves (30-)60-120(-150) cm long; petiole 10-30 cm long; 15-30 mm
diam., margins dentate; stipellae 5-8 mm long, narrow-lanceolate to lanceolate,
sometimes divided and 2-lobed from base, margins dentate, withering and appearing
spinose on older leaves; rachis slightly winged adaxially, wing continuous or as
separated segments, 1-3 mm wide, segments usually absent from petiole; pinnae
(7-)9-19; terminal pinna obovate and tapering into petiolule, lateral pinnae lanceolate
to oblong or to oblong-ovate or oblong-obovate with basal pair of pinnae broad-ovate
to ovate; apex acute to shortly acuminate; base rounded and ± equal in upper lateral
pinnae, unequal in basal pinnae; margins secondarily serrate, each tooth with a hair¬
like point; upper surface dark green and slightly glossy, sparsely hirsute, lower surface
fawn-hirsute; secondary veins 15-27 pairs, prominent on both surfaces; largest pinnae
(terminal and subterminal) (12-)18-35(-46) cm long, (6-)8-12(-16) cm wide; petiolules
0-5 mm long, up to 40 mm long on terminal pinnae. Inflorescences mostly cauliflorous
or ramiflorous, rarely borne amongst the leaves, pendent, 12-80 cm long; peduncle
(3-)10-18(-30) cm long; number of lateral spikes 2-24, 2-20 cm long; pedicels 1-5 mm
long; bracts amplexicaul, c. 5 mm long; bracteoles sessile, 3-5 mm long. Flowers 4- or
5-merous; sepals 5.0-8.0 mm long, recurved, basally connate, dark pink, outer surface
hirsute, inner surface pubescent, persistent in fruit; stamens 4.5-7.0 mm long,
filaments 3.5-4.6 mm long, anthers oblong, 2.0-3.0 mm long, yellow; ovary 2.3-3.0 mm
long, hirsute; styles 2 (or 3), divided almost to base, 5.0-8.0 mm long, glabrous.
Harden and Williams, Davidsonia (Cunoniaceae)
417
Fruit obovoid or ellipsoid, laterally compressed, 38-55 mm long, 32-53 mm wide,
35-46 mm deep; blue-black, glaucous, indumentum sparse at maturity (hairs visible
with hand-lens); mesocarp dark red; pyrenes usually 2, strongly laterally compressed,
3.0-6.0 mm long, fimbriate with radiating soft fibres attached to the pyrene body;
pyrene body 1.5-2.3 cm long, 1.5-2.1 cm wide, crest 5.0-10.0 mm long; seed ovate to
broad-ovate, 1.7-1.8 cm long, 1.2-1.4 cm wide, 3-4 mm deep. (Fig. 1).
Habitat and distribution: widespread in tropical rainforest in north-eastern
Queensland, chiefly from Cardwell area to Cooktown and inland to near Atherton
(Fig. 2). It grows from near sea level to altitudes of over 1000 m. Bange (1952)
erroneously cited this taxon as occurring in New South Wales. This taxon has been
collected in Fiji (Batiki, Naduruloulou) in 1949 by B. Parham (Parham 1972:125) where
it was cultivated at the Agricultural Experimental Garden (Bange 1952).
Phenology: chiefly flowers February to July, but recorded most months; with fruit ripe
mostly March to June, but recorded throughout the year.
Conservation status: due to its widespread distribution it is not considered to be
under threat; it is known to occur in a number of conservation areas, e.g. Lake Eacham
National Park, Bellenden Ker National Park (now part of Wooroonooran National Park).
Discussion: this species is widely cultivated as an ornamental tree and for its edible
fruits. As early as the 19th century (Bailey 1895,1898) this species was recommended
for cultivation because of its edible fruits.
Discussion of lectotype: the lectotype was designated by Bange (1952: 296) as
Murray's River, Rockingham Bay: Dallachy, 12 Aug 1866, a MEL collection with flowers
and fruit. There is no specimen at MEL that matches the original citation of Mueller;
however, there are nine sheets at MEL in type folders collected by Dallachy at
Rockingham Bay in 1866. Of these nine sheets only one has been annotated by Bange
(MEL 106575); this sheet has the upper portion of a leaf and fruit with a National
Herbarium of Victoria label, annotated 'Fragment of Type' with notes of Dallachy's
copied from MEL 106543. This sheet also has the following note: 'Dallachy made more
than one collection of this species, as these five sheets of types housed here include
flowers, fruits & foliage all with similar labels'. This label was not an original label and
the notes appear to have been copied in recent years (perhaps before the specimen
was seen by Bange in 1952). It also has a small label by Mueller with the name
Davidsonia pruriens.
The specimen (MEL 106543) which has the original of the notes on the habit of the
plant by Dallachy, including the date of 12 August 1866, also has a Phytologic Museum
of Melbourne label in Mueller's hand with 'Davidsonia pruriens Ferd. von Mueller,
Murray's River at Rockinghams Bay, Dali.'; the sheet has more recently been labelled
'Sheet 1 of 2'; this specimen is of the lower part of a leaf. Sheet 2 of 2 (MEL 106542)
has the upper part of a leaf and a Botanical Museum of Melbourne label in
Mueller's hand.
It is apparent that the specimens that were cited by Bange in 1952 have since been
remounted and possibly rearranged so that the specimen designated as the lectotype
(flowers and fruit) can no longer be identified as the one nominated by Bange. MEL
106546 is the only specimen that has an inflorescence and fruit and a label 'Davidsonia
pruriens' in Mueller's writing and a later National Herbarium of Victoria label with the
species name, locality as 'Rockingham Bay, Q'land', collector as J. Dallachy and date as
'71866'. We choose not to nominate this specimen as the lectotype; instead we choose
418
Telopea 8(4): 2000
Fig. 1. D. pruriens. a, adult leaf (x 0.1); b, stipules (x 0.5); c ( rachis, pinnae bases, stipellae (x 1.5);
d, inflorescence, diagramatic (x 0.1); e, part inflorescence (x 0.5); f, stamens (x 8); g, flower in L.S.
(x 3); h, flower (x 3); i, drupe (x 0.5): j, drupe in L.S. (x 0.5); k, pyrene (x 1); 1, pyrene in L.S. (x 1);
m, seed (x 1); n, embryo (x 1); o, embryo showing 2 cotyledons (x 1).
Harden and Williams, Davidsonia (Cunoniaceae)
419
to restrict the lectotypification to a single specimen, MEL 106575. This is in accordance
with the proposed amendment to Article 9 of the Code (Brummit 1998) which was
approved, in a slightly modified form, by the XVI International Botanical Congress.
This specimen was annotated by Bange, it consists of portion of a leaf and fruit and it
has a copy of the original notes of Dallachy with the date 12 th August 1866.
Common names: names used for this taxon include Davidsonian Plum by Bailey
(1898), Ooray by the Tully River aborigines (Bailey 1900), Davidson's Plum
(Mabberley 1997).
Selected specimens (from c. 36 examined): Queensland: Cook: Endeavour River, Persieh s.n., 1884
(MEL 106561); Bloomfield River, Barnard s.n. (MEL 106570); Daintree River, Pentzke s.n.,1882
(MEL 106557); near Lake Barrine, 10 miles [16 km] E. of Atherton, Melville 3683, Pont & Stephens,
9 Apr 1953 (NSW, BRI, K, MEL); Lake Eacham, Atherton Tableland, 17°17’S, 145°37’E, Williams s.n.,
29 Oct. 1987 (NE 61704); Copper Lode Falls Dam Site, on Freshwater Creek, approx. 6 miles
[9.5 km] S of Cairns, Cittins 2139, 22 Aug 1970 (BRI); Mt Bellenden-Ker, Mueller s.n., 1893
(NSW 104896, MEL); State Forest Reserve 185, Danbulla, Doggrell s.n., 28 Sept 1929 (BRI 333062,
L). North Kennedy: Tully, margin of Mitchell Park by stream, Clemens s.n., Dec 1949 (BRI 134733)
Fig. 2. Distribution of D. pruriens (A).
420
Telopea 8(4): 2000
2. Davidsonia jerseyana (F. Muell. ex F.M. Bailey) G. Harden & J.B. Williams, comb,
et stat. nov.
Davidsonia pruriens F. Muell. var. jerseyana F. Muell. ex F.M. Bailey, Queensland Flora 2:
538; Bange (1952: 296).
Type citation: Towards Tweed River'
Lectotype (Bange 1952: 296): New South Wales: North Coast: Richmond River;
Baeuerlen s.n., Oct 1892 (BRI11134).
Illustrations: Bange (1952: 295); Williams & Harden (1979:15); Harden (1990: 523).
Slender small trees to 6(-10) m high, unbranched or few-branched or with several
stems from the base, each bearing a terminal tuft of leaves; bark brown, corky and
somewhat scaly. Indumentum persistent on most parts, dense on young fruit and
shoots, hairs erect, pale brown, irritant; glandular hairs present on margins of stipules.
Seedling leaves at first simple, margins dentate and becoming Iobed at base, becoming
pinnate after 12 months growth, hirsute, with some reddish glandular hairs present on
stipules. Adult leaves (25-)35-80(-120) cm long; petiole 5-20 cm long; rachis
prominently winged adaxially, wing continuous or as separated segments, (l-)3-4
(-5) mm wide, becoming more separated on lower rachis as stipella-like outgrowths,
similarly on petiole, margins irregularly dentate, teeth ± spinose; pinnae (7-)ll-17
(-19); stipules, 10-30 mm diam., margins dentate; terminal pinna obovate to oblong-
obovate to elliptic, lateral pinnae mostly oblanceolate to oblong-oblanceolate, with
basal pinnae ± elliptic, apex acute to acuminate, base tapered and ± equal in upper
pinnae, unequal in lower pinnae, upper surface dark green and dull, sparsely hirsute,
lower surface dull and ± hirsute; margins secondarily serrate, each tooth with a hair¬
like point; secondary veins 11-17 pairs, prominently raised on both surfaces; terminal
and subterminal pinnae largest, (6-)20-30H0) cm long, lower pinnae (3-)6-10(-15.5) cm
wide; petiolules 1-3 mm long; stipellae present at base of most pinnae, lanceolate,
3—11 mm long, 3—5 mm wide, lower stipellae often 3-lobed, margins glandular¬
toothed. Inflorescences usually cauliflorous, mostly lateral on main stems, pendent,
compressed panicles, often appearing as tight dusters; 4-10(-30) cm long, peduncle
3-6 cm long, pedicels 1-3 mm long; bracts amplexicaul, 5.0-7.0 mm long, bracteoles
sessile 3.0-4.0 mm long. Flowers 4- or 5-merous; sepals 4-6 mm long, basally connate,
dark pink to red, outer surface bristly-hirsute, inner surface pubescent, recurved,
persistent in fruit; stamens 10, 4.5-6.5 mm long; filaments 4.1-5.5 mm long, anthers
ellipsoid, 1.1-1.7 mm long, yellow; ovary 1.5-2.6 mm long, styles usually 2, divided to
base, 4.4—5.6 mm long, glabrous; stigmas minutely capitate. Fruit pyriform to obovoid,
laterally compressed, 33-45 mm long, 31-37 mm wide, 27-35 mm deep, blue to black,
glaucous; young fruits hirsute, mature fruits sparsely hirsute with golden-brown to
fawn hairs 1.0-2.0 mm long; mesocarp dark red; pyrenes usually 2, strongly laterally
compressed, fimbriate with soft fibres 1.0-2.0 mm long, attached to the pyrene body;
pyrene body 1.4-1.8 cm long, 0.9-1.5 cm wide, crest 2.1-2.3 mm long; seed 1.1-1.2 cm
long, 0.8-0.9 cm wide, 2.5-3.0 mm deep. (Fig. 3).
Habitat and distribution: usually grows in subtropical and riverine rainforest, in
moderately high rainfall from 1100 to over 2000 mm, at altitudes less than 300 m above
sea level. Mostly on red and yellow podsolic soils of clay texture, over fine-grained
metasediments, greywacke, slate, phyllite or quartzite, also on alluvial deposits.
Confined to north-eastern New South Wales; chiefly from the Brunswick and Tweed
Rivers catchments, from Mullumbimby north to Urliup, Upper Crystal Creek and
Settlement Road (Fig. 4).
Harden and Williams, Davidsonia (Cunoniaceae)
421
Fig. 3. D. jerseynna. a, adult leaf (x 0.2); b, stipules (x 0.5); c, rachis, pinnae bases, stipellae (x 1.5);
d, inflorescence (x 0.5); e, part inflorescence (x 1); f, stamens (x 8); g, flower in L.S. (x 3); h, flower
(x 3); i, drupe (x 0.5): j, drupe in L.S. (x 0.5); k, pyrene (x 1); 1, pyrene in L.S. (x 1).
422
Telopea 8(4): 2000
Phenology: flowers recorded chiefly from October to January; fruit ripe mostly
December to February.
Conservation status: listed in New South Wales as an endangered species on Schedule
1 of the Threatened Species Conservation Act, 1995 ; 2ECi on the ROTAP List (Briggs &
Leigh 1996). D. jerseyana is only known to occur in one gazetted reserve, Brunswick
Heads Nature Reserve. The largest known population of this taxon is just to the west
of the Brunswick Heads Nature Reserve (Watson 1987). Many of the smaller
populations are on the verges of minor roads; several are represented by isolated trees,
and therefore may not be viable in the longer term.
Discussion of type: there is no description of this taxon by Mueller; but, 'in an
unpublished, and apparently (Ross, pers. comm.) now no longer extant manuscript, he
named this new form D. jerseyana’ (Watson 1987 p. 4). However, Hoogland in 1978
examined MEL specimens 'Taken from a folder marked Geissois jervoisiana F. Muell.
ms.' which he determined as Davidsonia pruriens. The specimens annotated by
Fig. 4. Distribution of D. jerseyana (V) and D. johnsonii (•).
Harden and Williams, Davidsonia (Cunoniaceae)
423
Hoogland were Baeuerlen 864 (MEL 106568 and MEL 106569) and Barnard s.n. (MEL
106570 and MEL 106571). The former specimens have since been re-determined as
D. jerseyana and the latter confirmed as D. pruriens. MEL 106568 has attached a draft
description of the specimen in Mueller's hand, but, there is no annotation of the name
on either the note or on the sheets by Mueller.
The first reference to this new taxon was in Bailey (1895), 'D. jerseyana, F. v. M., found
on the southern border of Queensland and in the adjoining scrub lands of New South
Wales'. Bailey (1900) listed var. jerseyana (with D. jerseyana F. v. M. as a synonym) as a
variety, validating the name with the brief note that it is smaller in all parts than the
type variety. It was not till 1909 that Bailey (p. 169) gave his own name as author of the
variety, when he listed under Davidsonia, var. jerseyana Bail. = D. jerseyana F. v. M.
As no holotype existed, Bange (1952) proposed the specimen collected by Baeuerlen in
October 1892 (BRI 11134) at Richmond River (New South Wales) as the lectotype for
the variety 'because it is probably the specimen F. v. Muell. and Bailey have seen, the
more so as the specimen is a good one'. Bange made no mention of the specimens
collected in the same month in 1892 by Baeuerlen (No. 864) at Murwillumbah (MEL,
NSW) as those at MEL were under the genus Geissois and he did not examine NSW
material. It could be argued that the locality (Richmond River) of the designated
lectotype does not fit Bailey's published habitat description of 'Towards Tweed River .
However, Bailey gave that vague description in The Queensland Flora (1900) from a
Queensland perspective and we do not consider it sufficient reason to overturn
Bange's lectotypification. This taxon is not known to occur in Queensland, even
though it has been found very close to the border with that State; Bange (op. cit.)
erroneously cited a specimen collected at Chillingham ( Dixon s.n., BRI 101933) for
Queensland.
Common name: the common name widely used for this taxon is Davidson's Plum
(Williams et al. 1984; Floyd 1989; Harden 1990).
Selected specimens (from 14 examined): New South Wales: North Coast: Chillingham Road, 13 km
NW of Murwillumbah, Foreman & Woodland s.n., 22 Oct 1977 (NE 35399); Chillingham, Tweed River,
Dixon s.n., Oct 1933 (BRI 333069); Murwillumbah, Baeuerlen 864, Oct 1892 (NSW, MEL); Tweed
River, Fawcett s.n., 1883 (MEL 106559); 5 km NE of Mullumbimby, Eliott s.n., 18 Aug 1991 (NE
55496); 1km E of Mooball, Williams & Harden s.n., 17 Nov 1979 (NE 61638, NSW, BRI, CANB);
1 km east of Mooball, beside Burringbar Creek, Williams s.n., Nov 1980 (NSW 389178, MEL, NE);
Brunswick River, White s.n., Nov 1885 (MEL 106549); Brunswick River Bridge, Floyd & Hayes s.n.,
21 Jan 1958 (CFSHB).
3. Davidsonia johnsonii j.B. Williams & G. Harden, sp. nov.
Ab aliis speciebus combinatione characterum sequentium distinguitur: planta in
omnes partes plus minusve glabra; folia minora pinnis paucioribus; inflorescentiae
terminates vel subterminales in ramulis frondosis; fructus depresse globosus;
indumentum calycis breviter molliterque pubescens; pili irritantes omnino deficientes.
Type: New South Wales: North Coast: Huonbrook, Wilsons Creek Road, west of
Mullumbimby, J.B. Williams & G.J. Harden s.n., 17 Nov 1979 (holo NSW 427612; iso AD,
BRI, CANB, HO, K, MEL, NE)
[Davidsonia species in Williams & Harden (1979:15); Williams et al. (1984: 30)]
[Davidsonia species A in Harden (1990: 523)]
[Davidsonia sp. A Mullumbimby-Currimbin [sic] Ck (A.G. Floyd 1595) in Threatened
Species Conservation Act, 1995]
424
Telopea 8(4): 2000
Illustrations: Williams & Harden (1979: 15); Harden (1990: 523).
Bushy spreading, suckering small tree 5-10(-18) m high, well-branched and forming
a dense leafy crown; freely suckering from the roots; bark fawnish, smooth or
becoming finely scaly. Indumentum inconspicuous or absent on mature parts; buds,
young shoots and leaves pubescent, glabrescent with age; inflorescence axes and
perianth shortly and finely pubescent; leaf buds fawn pubescent with non-irritant
hairs; stalked red glandular hairs present on young shoots, and terminating teeth on
rachis wings, margins of stipules and pinnae. Seedling leaves not seen as no viable seeds
are known to develop. Adult leaves ; (6-)10-33(-47.5) cm long; petiole 2.5-7.0 cm long;
rachis prominently winged adaxially, wing mostly continuous on rachis, as separated
segments on petiole, wing irregularly dentate, each tooth with a hair-like point;
stipules 7-15 mm diam., margins dentate, persistent; pinnae (3-)5-9(-ll); terminal
pinna mostly obovate to oblanceolate and tapering into petiolule, lateral pinnae
narrow-ovate to narrow-obovate to oblong-oblanceolate, basal pinnae ovate to falcate-
elliptic; apex acute to slightly acuminate, base ± equal in upper pinnae, unequal on
lower pinnae, upper surface green and glossy, lower surface dull, and both surfaces
glabrous, margins regularly dentate; secondary veins 6-9 pairs, raised on lower
surface; largest pinnae (terminal or subterminal) usually (10-)13-17(-25) cm long,
(4-)5-7(-8) cm wide; lower pinnae 1.7-9.3 cm long, 0.8-4.5 cm wide; petiolules 0-2 mm
long, except for terminal petiolule 10-25 mm long; stipellae usually present at base of
pinnae (especially lower pairs), narrow-lanceolate, 3-11 (-20) mm long, 1-3 mm wide,
margins glandular-toothed. Inflorescences borne amongst the leaves, often terminal,
elongated panicles, usually 10-20 cm long; peduncle 0.5-5 cm long; pedicels 0-1.0 mm
long; bracts 3.0-4.0 mm long; bracteoles sessile, 2.0-3.0 mm long. Floivers 4- or 5-
merous; sepals 3.1 —4.4 mm long, dark pink, both surfaces finely and shortly pubescent;
stamens (6-)8-10, 3.5-5 mm long; filaments 4.1-6.0 mm long, anthers ellipsoid to
obovoid, 1.4-2.2 mm long, yellow, pollen absent from flowers examined; ovary 1.0-1.7 mm
long, pubescent, styles 2 or 3, 2.1—4.5 mm long, often reduced or absent; sometimes
ovary vestigial or lacking. Fruit depressed-globose, 20-39 mm long, 25-60 mm wide,
28-53 mm deep, reddish purple to purplish black; sprinkled with fine hairs, appearing
smooth; mesocarp reddish; pyrenes usually 2 per fruit, sometimes 3, shape variable
dependent on number, surface shortly and softly fibrous; pyrene body 0.9-1.2 cm long,
1-0—1 • 1 cm wide, crest 1.4-1.7 mm long; seeds not known to develop within the
pyrenes. (Fig. 5).
Habitat and distribution: Davidsonia johnsonii usually grows in disturbed subtropical
rainforest, or on margins of wet sclerophyll forest and gully rainforest, sometimes in
disturbed areas, mostly from 15 to 260 m altitude. Usually on podsolic soils of variable
structure, gleyed podsolics or kraznozems on basalt, often over Silurian greywacke,
slate or phyllite; over Lismore basalt and its boundary with Nimbin rhyolite; and over
Lamington volcanics. Dominant species include Eucalyptus grandis and Lophostemon
confertus in wet sclerophyll forest, and with many species in rainforest including
Hicksbeachia pinnatifolia, Diploglottis australis, Acacia melanoxylon, Castanospora alphandii,
Omalanthus populifolius.
This taxon has a very scattered distribution and is known from a limited number of
small populations, north from Tintenbar in New South Wales to near Natural Bridge
(Watson 1987) and Upper Tallebudgera (McDonald pers. comm. 2000) and Upper
Currumbin valleys in southern Queensland (Fig. 4), a distance of about 120 km and an
east-west range of 30 km. Mostly occurs as groups of several mature trees, with many
saplings and young plants that have developed from root suckers.
Harden and Williams, Davidsonia (Cunoniaceae)
425
Fig. 5. D. johnsonii. a, adult leaf (x 0.3); b, stipules (x 1.5); c, rachis, pinnae bases, stipellae (x 1.5);
d, inflorescence (x 0.5); e, stamens (x 8); f, flower in L.S. (x 3); g, flower (x 3); h, drupe (x 0.5):
i, drupe in L.S. (x 0.5); j, pyrene (x 1); k, pyrene in L.S. (x 1).
426
Telopea 8(4): 2000
Phenology: flowers chiefly October to November; ripe fruit recorded mostly February
to April.
Conservation Status: listed as 'Davidsonia sp A Mullumbimby-Currimbin [sic] Ck (A.G.
Floyd 1595)' in New South Wales as an endangered species on Schedule 1 of the
Threatened Species Conservation Act, 1995; 2ECi on the ROTAP list (Briggs & Leigh
1996).This species is only known to occur in one gazetted reserve. Snows Gully Nature
Reserve in New South Wales. Tire largest populations at Huonbrook and Upper
Burringbar are on private properties; present owners are actively involved in the
conservation and rehabilitation of disturbed rainforest. Threats to the species include
general clearing of remnant pockets of rainforest and the likely narrow genetic base in
a limited set of small populations.
There are some important questions arising as to the dispersal of the species and its
future in the wild. Flowers are formed regularly and prolifically, but as indicated in the
species description no pollen has been detected in mature anthers and ovaries are
small or lacking, sometimes vestigial and often the styles are reduced or absent.
Ovules are usually absent — but in one collection two or three abortive shrivelled
ovules were present in each of the two loculi. Fruit are produced abundantly and are
well-developed but the pyrenes are always empty and no seed has ever been found.
From our own field observations and communications with others that are familiar
with field populations (G. Watson, A.G. Floyd, D.L. Jones) the species relies on root
suckers for reproduction. This vegetative reproductive process readily enables present
stands to propagate, but it does not account for the species distribution in populations
scattered over 120 kms. Unless, as suggested by Watson (1987), aboriginal people were
involved with the vegetative spread of this species over its geographic range. Or,
alternatively is it possible that very rarely fertile seeds are formed, with subsequent
dispersal and establishment of seedlings.
Derivation of name: named in honour of the late Dr L.A.S. Johnson who during the
past 49 years made outstanding contributions to systematic botany, rainforest floristics
and to the understanding of the Australian flora while on the staff and as Director, and
latei as Director Emeritus, of the Royal Botanic Gardens Sydney. Lawrie Johnson
annotated specimens of this taxon in 1949, the year after he joined the staff of the
Gardens, as 'clearly a new species'.
Discussion: a specimen of this taxon collected in 1936 at Nimbin by King (BRI) was
cited by Bange (1952) as 'variety dubious'. Specimens deposited at Coffs Harbour
Forestry Herbarium (now CFSHB) collected by Floyd and Hayes at The Pocket in 1958
and Watson at Thieles Gate, Huonbrook in 1978 were originally not separated as a new
taxon. It was not until Watson brought the plants to the attention of the authors in 1978
that it was further investigated and the differences from the other two taxa were
clarified.
Common names: Smooth Davidsonia (Williams et al. 1984), Smooth Davidson's Plum
(Floyd 1989, Harden 1990), Small4eaved Davidson's Plum (Floyd 1989).
Selected specimens (from 21 examined): Queensland: Moreton: Olson's Property, Currumbin
Valley, Jones s.n., 10 Jan 1982 (NSW 389182, BRI, CANB, NE).
SctfuT, l !lu Wal t!;, NOrth C ° aSt: Ho § ans Sc* -1 *' North Tumbulgum, Floyd 1827, 8 Dec 1981 (NSW,
CFSHB); Whian Whian, Chee! s.n., 6 Sept 1926 (NSW 104890); Mooball, near Murwillumbah, Flowers
s „"’' 2 Ma y 1944 (NSW 104891); 5.2 km west of Burringbar on property of M. Lickfold, Williams,
mp pp 1980 (NSW 389176 ' A ' AD ' B ' BR1 ' CANB '™, HO. K, L, LE, MO,
G.,'‘ b ' 1 RE 'V S 'Burringbar Range, end of track 1.7 km N of Upper Burringbar, 22°25'S
153 27 E, Davws 1564a & Richardson, 27 Feb 1990 (CANB, NSW, BRI, MEL); Nimbin, King s.n., 2 Mar
Harden and Williams, Davidsonia (Cunoniaceae)
427
1936 (BRI333410); Kerle's Property, c. 9 km W of Mullumbimby on Wilsons Creek Road, Williams s
17 Dec 1985 (NE 69307, NSW, BRI, MEL); Thiele's Gate, Huonbrook via Mullumbimby, Watson s.n.,
26 Aug 1978 (NE 35678), Floyd 1595, 3 Dec 1980 (CHSHB); The Pocket, Billinudgel Rd, Floyd &
Hayes s.n., 22 Jan 1958 (CFSHB); Broken Head, 10 km S of Byron Bay, Eliott s.n., 25 Aug 1991
(NE 55495).
Acknowledgments
Our thanks to Graham Watson, Michael Lickfold and other conservationists who
assisted us in locating populations in the field, and to colleagues for commenting on
the manuscript. We are grateful to Nicola Oram and Lesley Elkan for the illustrations,
Wayne Cherry for generating the distribution maps. Dr Peter Wilson for the Latin
diagnosis and advice on the lectotypification of the species, and Kristina McColl and
Wayne Cherry for general assistance. The heads of the following herbaria are thanked
for allowing access to their collections either on loan or in situ: BRI, CFSHB, MEL, K,
NE, NSW, QRS. Thanks also to Nan Nicholson and Brett O'Donovan (Terania
Rainforest Nursery Pty Ltd) for providing seedling material.
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Manuscript accepted 22 March 2000
429
Seedling characteristics in the Casuarinaceae
Robin Hwang and John G. Conran
Abstract
Hwang, Robin ! and Conran, John G. 2 ’ 3 (IResearch Division, National Museum of Natural Science,
Taichung, Taiwan; 2 Department of Environmental Biology, The University of Adelaide, SA, Australia 5005;
3 author for correspondence) 2000. Seedling characteristics in the Casuarinaceae. Telopea 8(4): 429-439.
Seedling characters and their distributions within the genera of the Casuarinaceae and their
sections are described for 45 taxa in the family representing three of the four genera and all 14 of
the currently recognised sections in Allocasuarina. The cotyledons in Gymnostoma are petiolate, a
feature absent from the other taxa examined. There were no obvious genus-level seedling features
distinguishing either Casuarina or Allocasuarina, although red-purple cotyledons at senescence are
common in Casuarina but rare in Allocasuarina. Several features are informative at the sectional level:
suppression of secondary cotyledon branches is only found in Allocasuarina sects, Oxypitys,
Platypitys, Echinopitys and Ceropitys, as were abortion of the primary shoot and divergent tooth
apices, the latter two characters otherwise found only in A. monilifera and A. microstachya. Seedlings
of sect. Ceropitys showed a distinctive growth habit making them look like Lycopodium shoots with
outgrowths resembling microphylls.
Introduction
The Casuarinaceae are an unusual family of dicotyledons from SE Asia, Australia and
Oceania. Within the Casuarinaceae there are four genera and about 80 species of which
three genera and 66 species occur in Australia, most of which are endemic (Johnson
& Wilson 1993). Bentham (1873) classified the species largely on cone bracteoles
(valves), whereas Diels and Pritzel (1905) emphasised features of the vegetative
branchlets. Modern relationships in the family are based largely on Poisson's (1874)
division of Casuarina sens. lat. into the Gymnostomae (= Gymnostoma) and Cryptostomae
(the remainder) using stomatal features. Barlow (1959) divided the Cryptostomae into
'Group A' and 'Group B' based on cytology, and Johnson (1982) expanded Barlow's
ideas, establishing Allocasuarina for Group B with Group A representing Casuarina
s. str. The Gymnostomae were raised to generic rank as Gymnostoma (Johnson 1980) with
c. 18 species from SE Asian wet tropical forests, and Ceuthostoma for two Malesian
Gymtwstoma-like species which lack a broad bract under the bracteole pair and which
possess antrorse anthers and very long-bracteolate cones on long branchlets
(Johnson 1988).
There are three genera in Australia (Wilson & Johnson 1989): Gymnostoma has a single
extant native species, although fossil evidence suggests that the genus was much more
diverse during the Eocene (Christophel 1980, Scriven & Christophel 1990) and
Palaeocene (Scriven & Hill 1995). Casuarina has six widespread native species,
generally on fertile soils, whereas the endemic Allocasuarina with 14 sections and 60
species is frequently associated with nutrient-deficient soils (Wilson & Johnson 1989).
Although many Allocasuarina sections are mono- or oligotypic, most species belong to
sect. Cylindropitys (29 spp.) or sect. Ceropitys (nine spp.). Cylindropitys, in particular,
consists of many closely related taxa thought to represent a recent evolutionary
radiation (Wilson & Johnson 1989).
430
Telopea 8 ( 4 ): 2000
Cytology and karyotype (Barlow 1959,1983) support several of the fruit, bracteole and
samara-based subdivisions in the family (Johnson & Wilson 1989). There are also
studies detailing pollen (Kershaw 1970), anatomy (Williams & Metcalf 1985), stem
morphology (Torrey & Berg 1988), 17 stem, tooth and stomate characters in
Gymnostomci and the other genera in the family, especially as fossils (Scriven &
Christophel 1990, Scriven & Hill 1995), and 22 vegetative, stomatal and reproductive
characters for extant genera (Dilcher et al. 1990).
De Candolle (1846) stressed the taxonomic value of seedling morphology. Seedling
characters were deemed useful in angiosperm classification (de Vogel 1980), especially
if adult material was difficult to obtain or the plants had long life cycles. Clifford (1991)
and Li and Hsieh (1997) observed that members of the same genus often have similar
seedling characteristics. Leonard (1957) advocated that there should be only one type
of seedling in a 'good' genus, and that genera should differ significantly in their
seedling morphology. Nevertheless, Weberling and Leenhouts (1966) proposed that
genera should not necessarily be divided on seedling variation. Conran et al. (1997)
found that for some taxa, seedling characters were more useful at subgeneric and
sectional levels, rather than generic.
Boodle and Worsdell (1894), commenting on the higher level relationships of
Casmrim, included seedling information, and Duke (1965), Burger (1972), Torrey
(1983) and Boland et al. (1984) described or illustrated seedling morphology for a
number of Casuarinaceae species. Their listed characteristics included root colour;
lateral root abundance; hypocotyl position against the soil; hypocotyl colour and
length; cotyledon shape, index, attachment, apex and base shape, adaxial and abaxial
colour, texture and indumentum; epicotyl length (first internode above the hypocotyl),
stem straightness; number of leaves (teeth) per whorl and tooth shape, colour and
length. Unfortunately data were provided for few taxa, and generally without
systematic sampling within the family.
This present study investigates seedling morphology in the Casuarinaceae, comparing
the results against the current classification of Wilson and Johnson (1989) and the adult
morphology-based studies of Dilcher et al. (1990) and Scriven and Hill (1995) to
examine the distribution and usefulness of seedling features in the family.
Materials and methods
Seeds from 123 provenances representing eight Casmrim and 34 Allocasuarina taxa
were obtained from various sources detailed previously in Hwang and Conran (1991).
The seeds were soaked overnight under running tap water, and germinated on wet
cotton wool in an unheated glass house (temperature range 10-35°C) under natural
lighting. Casuarinaceae seeds germinate readily, mostly in 15-30 days at 20-25°C
(Elliot & Jones 1982), and most species benefit from daily exposure to light during
germination (Turnbull & Martensz 1982). Pre-soaking the seeds in water for 24 hours
improves germination rates (Kuo 1984). A summary of the germination features for
Australian Casmrim and Allocasuarina species is given in Hwang and Conran (1991).
Germinated seeds were planted 40 mm apart and 10 mm deep in drained plastic trays
with an unfertilised, unsterilised 2:2:1:1 soil mixture of coarse river sand, fine pine
bark, red mountain soil and grey sandy loam. There were 15 columns x 7 rows per tray,
with 21 seedlings per provenance. Average seedling survival per provenance was 18.8
(range 1 to 21), and the seedlings were grown and observed for 3 months with daily
watering. With the exception of ephemeral or developmental characters, data were
recorded from three month old seedlings and scored as the average of all surviving
seedlings for each taxon. Data for Casmrim junghuhniana Miq. were taken from the
Hwang and Conran, Seedling characteristics (Casuarinaceae)
431
published description of its seedlings by Burger (1972). Seedling data for Gymnostoma
australianum L. Johnson were obtained from plants grown from seeds collected at
Noah Creek (Qld) by Dr D.C. Christophel and used in a study by Prider (1998).
The seedling characters and their states were recorded as follows:
(1) Primary roots red (1) or white to yellowish (0).
(2) Cotyledon orientation in relation to the stem apex (Fig. la): level (0), ascending (1)
or descending (2).
(3) Cotyledon apex shape (Fig. lb): rounded (0), obtuse (1), acute (2).
(4) Base of the cotyledons (Fig. lc): not markedly constricted (0), constricted (1),
petiolate (2).
(5) Cotyledon shape in cross-section (Fig. Id): flat (0), slightly convex (1), convex (2).
(6) Colour of senescing cotyledons: yellow to light brown (0) red purple to dark rust-
red (1).
(7) Cotyledons pubescent: absent (0), present (1).
(8) Number of axillary branchlets developing in the cotyledons.
(9) Cotyledonary axillary branchlets: only developing after those of the primary stem
(1) ; developing at the same time as the primary stem (0).
(10) Epicotyl-base colour: green to yellow (0) or red to purple (1).
(11) Primary shoot orientation (Fig. le): erect (0), slightly curved (1), strongly curved
(2) or recurved/curled (3).
(12) Primary branching (Fig. If): simple (0) or compound (1).
(13) Node number on the main shoot at which branching first occurs (Fig. lg)
excluding the cotyledonary node.
(14) Axillary branch supression zone (Fig. lh): absent (0); present (1).
(15) Primary stem: developing normally (0); suppressed and replaced by cotyledonary
or basal lateral shoots (1).
(16) Lateral branchlets (Fig. li): strongly curved (1); or straight (0).
(17) Number of teeth primary shoot node 15.
(18) Tooth bases on lateral branchlets (Fig. lj): imbricate (1); valvate (0).
(19) Lateral branch tooth apices (Fig. Ik): closely appressed (0); or with loosely
attached, spreading, or divergent apices (1).
(20) Seedling growth form: stems jointed but without obvious microphyll-like
protrusions (0); strongly resembling Lycopodium shoots and having the appearance of
bearing microphylls (1).
Results
The results characters are summarised in Table 1.
The primary root (character 1) was red in Gymnostoma and the majority of Casuarina
and Allocasuarina sects Allocasuarim, Cylindropitys, Nannopitys, and Trichopitys, while
Allocasuarina sects Dolichopitys, Echinopitys and Oxypitys had yellow- or white-rooted
seedlings although both colour types were represented in most sections.
The cotyledon orientation (2) was horizontal in the majority of the taxa examined,
except for A. acutivalvis and A. paradoxa where it was descending, and A. thuyoides
where it was ascending. Tire cotyledon apex (3) was usually rounded, although it was
obtuse in four species of Allocasuarina, mostly in sect. Cylindropitys, and acute in four
other species of Allocasuarina. Cotyledon bases (4) were distinctly petiolate only in
Gymnostoma, not only for G. australianum, but also in seedlings of G. vitiense which
were grown for this study but which did not survive beyond the cotyledon stage. The
base of the cotyledons was markedly constricted in all Casuarina spp., but four species
of Allocasuarina in different sections. Cotyledon shape in cross-section (5), although
432
Telopea 8 ( 4 ): 2000
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Fig. 1. Seedling characteristics of the Casuarinaceae, details outlined in the materials and methods,
a, cotyledon orientation; b, cotyledon apex shape; c, cotyledon basal constriction; d, cotyledon
cross-sectional shape; e, shoot orientation (bending); f, primary branches simple or compound;
g, node number at first branching; h, axillary branch sterile zone; i, lateral branch curvature;
j, lateral branch tooth bases; k, lateral branch tooth apical spread. Character code numbers are those
listed in the text.
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Taxon Character Number
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Telopea 8(4): 2000
constant within species, was highly variable within the genera and sections, with no
clear patterns apparent.
The colour of the cotyledons at senescence (6) differed from the pattern seen for roots
of the same species, and was divided into two states: yellow to light brown versus red-
purple to dark rust-red, the latter was the most common condition in Gymnostomn and
Casuarina, and yellow or light brown in Allocasuarina. Cotyledon pubescence (7) was
uncommon, but was observed in three Allocasuarina species in different sections.
In A. acutivalvis, A. campestris, A. decaisneana and A. tessellata, the interpetiolar
branchlets (9) always developed before those of the cotyledons, and these were then
followed by lateral branchlets on the main shoot. In contrast, only interpetiolar
branchlets were seen in C. cunninghamiana, C. cristata, C. obesa and C. glaitca and then
only developing later than and posterior to the lateral branchlets. In 12 species, apical
dominance of the primary shoot was suppressed, with replacement bv axillarv
branchlets (15). y y
Epicotyl-base colour (10) varied from green or yellow through to red or purple. As
with cotyledon senescence, red or purple was most common in Casuarina. A. torulosa
was the only Allocasuarina with a red-purple epicotyl and cotyledons.
Primary shoot shape (11) varied from erect (Gym nostoma, Casuarina and some
Allocasuarina spp.), slightly curved (various Allocasuarina spp.), to strongly curved (the
predominant condition in sections Echinopitys, Ceropitys) or even curled (A. scleroclada
and A. striata). The lateral branches were also strongly curved in three species of
Allocasuarina including some, but not all, species where the primary shoot curled or
was strongly curved. The primary branches (12) were generally compound, but simple
Tranches occur in a number of species, although there were no clear supra-specific
patterns. Nevertheless, simple cotyledon branches were only found in Allocasuarina
sects Dolichopitys, Oxypitys, Platypitys, Echinopitys and Ceropitys. The node number on
the mam shoot at which branching first occurred (13) was also largely constant within
but variable between, species ranging from the second node in A. paradoxa to the
twelfth node in A lehmanmam (both sect. Cylindropitys). There was an axillary branch
sterile zone (14) present in C. cristata and A. thuyoides. In sections Oxypitys, Platmitys
Echinopitys almost all of sect. Ceropitys, and A. monilifera and A. microstachya, a) the
primary shoot aborts (15), and the main adult stem or stems develop from axillarv
shoots arising from the cotyledon axils, and b) the lateral branch stem tooth apices
were divergent. r
Node 15 on the primary stem (17) in Gymnostoma was 4-toothed, with the remainder
u u, TT 6S 6X , a ? lined bein 8 4 ‘ and 6-toothed (rarely 5-toothed). On lateral
branchlets the tooth bases (18) were imbricate in Allocasuarina sect. Oxypitys Seedlings
shoots°ofthp C r° pih J S : U P toabout three months old, superficially resemble
shoots of the fern ally Lycopodium, where there are protrusions along the stems which
give the impression of micophylls — a feature also noted by Torrey (1983).
Discussion
Seedling structure within the family is distinctive and highly derived, reflecting the
unusual adult morphology. Accordingly, meaningful comparisons with the seedlings
different fl a ^ eS & r ^ data revealed * number of patterns inlhe
single cha f rarlL reS 'f G r M °i m ' 7 Se ? dM % s ' 111 P^ular, are readily identified by the
s ngle character of strongly peholate cotyledons. Casuarina seedlings are not clearlv
istinguished from those of Allocasuarina, although they tend hAave constricted
cotyledon bases, and the senescent cotyledons and epicotyl are generS^rTto
Hwang and Conran, Seedling characteristics (Casuarinaceae)
437
purple. Dilcher et al. (1990) and Scriven and Hill (1995) similarly found that both
extant and fossil Casuarinaceae can be identified and assigned to genera on the basis
of vegetative and reproductive anatomy. Red roots in the Casuarinaceae and some
other families indicate high levels of leghaemoglobin (Martin & Dowd 1993), where it
is linked to nitrogen fixation by the soil actinomycete Frankia (Maggia & Bousquet
1994). Although this is common throughout the family, it is mixed enough in its
occurrence within each of the genera and sections to suggest that the feature has
limited phylogenetic utility within the family.
In the Casuarinaceae, there are also several taxa where the seedling characteristics
differ from those of the adults. For example, lateral branch tooth apices are divergent
in juveniles of sect. Oxypitys, but appressed in the adults, and appressed in seedling
A. dielsiana, A. Iieugeliana and A. pusilla (amongst others), but divergent in the adults.
Similarly, whereas no Casuarina species have divergent teeth in the seedlings, adult
C. obesa and C. glauca both possess them, hr A. acutivalvis the branchlets changed
gradually into the adult form, whereas in A. campestris and A. tessellata one or a few
axillary shoots develop directly with adult morphology. Torrey (1983) also observed
that adult branchlets develop after about 10 cm of juvenile growth, but in A. acutivalvis
there can be reversal, where adult shoots revert to the juvenile form.
A phenetic analysis of the Casuarinaceae by Hwang (1989) using the characters
described here, plus a series of growth rate-related characters found that there were
several distinctive species assemblages, representing in decreasing order of
dissimilarity: Allocasuarina sect. Oxypitys-, a mixed group from Allocasuarina sects
Ceropitys and Echinopitys; Casuarina sens. str. plus Allocasuarina torulosa (Aiton)
L.Johnson; and then three groups consisting of mixtures of taxa from the remaining
Allocasuarina sections sampled (see taxa listed in Hwang & Conran 1991).
Prider's (1998) viability and embryological studies of Gymnostoma australianum
demonstrated that the majority of the samaras produced were embryologically non-
viable rather than short-lived, and that some of the few fertile seeds produced were
still viable after 2 years. Similarly, Torrey (1983) was uncertain whether germination
failure in several of the species that he examined was due to seed age or general low
viability.
Relationships between series and sections in large genera can often be defined, at least
in part, on seedling structure. Conran et al. (1997) found distinctive seedling-based
groups in the Droseraceae which are taxonomically informative at the subgeneric and
sectional levels, and which indicated that there are previously unpredicted
relationships between them. Within Acacia (Mimosaceae), seedlings are distinctive at
the subgeneric level (e.g. Burger 1972) and reflect features which, though present in
the seedlings, are absent in mature plants. In Eucalyptus (Myrtaceae) the seedlings are
also different between subgenera and between some of the sections (Chippendale
1988), with Maiden (1929-31) dividing the genus into three sections on cotyledon
shape. Similarly, the E.flocktoniae and E. transcontinentalis complexes of series Subulatae
both possess uniquely decussate-leaved seedlings (Nicolle & Conran 1999). Weberling
and Leenhouts (1966), Burger (1972) and Li and Hsieh (1997) also noted many
examples of variable and potentially taxonomically useful seedling characters at the
intrafamilial and infrageneric levels in a range of rainforest taxa.
The patterns seen here for some of the seedling characteristics in the Casuarinaceae
suggest that there are some distinctive seedling-based species groupings. Allocasuarina
sect. Ceropitys have lycopod-like seedlings, and simple cotyledon branches.
Allocasuarina sects Dolichopitys, Oxypitys, Platypitys, Echinopitys and Ceropitys, had
abortion of the primary shoot and divergent tooth apices characters otherwise found
only in A. monilifera (sect. Cylindropitys) and A. microsiachya (sect. Trachypitys).
438
Telopea 8(4): 2000
Although the treatment of the Casuarinaceae by Wilson and Johnson (1989) placed
taxa close to their presumed relatives, there is no current phylogenetic classification of
the family from which species relationships can be investigated. Our study provides
good evidence that there are useful seedling characteristics in the Casuarinaceae
which might be used to investigate generic, sectional and species relationships within
the family. However, it is not the intention of this paper to reject one set of characters
used to classify the family in favour of another. Rather, it is hoped that the seedling
characters presented here can be incorporated with as wide as possible a range of
features from adult, seed and pollen morphology plus cytology so that future
phylogenetic studies of the family can assess objectively the relative contribution and
merits of all the available information.
Acknowledgments
This work was undertaken as part of a Ph.D. by RH, and the Department of Ecology
and Evolutionary Biology at Monash University is thanked for the provision of
facilities, as is the Department of Environmental Biology at the University of Adelaide
for supporting RH as a distinguished visiting fellow. The late Dr L.A.S. Johnson is
thanked for stimulating comments on this research.
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Clifford, H.T. (1991) Germination patterns in Dicotyledons. Aliso 13: 207-213.
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de Vogel E.F. (1980) Morphological types in dicot seedlings with reference to their origin. Bull Soc
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Dilcher, D.L., Christophel, D.C., Bhagwandin, H.O., Jr. & Scriven, L.J. (1990) Evolution of the
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EHiot, W.R. & Jones, D.L. (1982) Encyclopaedia of Australian Plants suitable for Cultivation vol 2
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Hwang, Y.H. (1989 ) ) Taxonomy of Australian Casuarinaceae based on seedling morphology and allozymes
Unpublished Ph.D. thesis. Monash University. 5 P ^ uuuzymes.
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Hwang, Y.H. & Conran, J.G. (1991) Seedlings of Australian Casuarinas. I: Germination. IV. Austral.
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Johnson, L.A.S. (1980) Notes on Casuarinaceae. Tclopca 2: 83-84.
Johnson, L.A.S. (1982) Notes on Casuarinaceae II. J. Adelaide Bat. Card. 6: 73-87.
Johnson, L.A.S. (1988) Notes on Casuarinaceae Ill: The new genus Ceuthostoma. Telopea 3:133-137.
Johnson, L.A.S. & Wilson, K.L. (1989) Casuarinaceae: a synopsis. Pp. 167-188, in Crane, P.R. &
Blackmore, S. (eds). Evolution, Systematics and Fossil History of the Hamamelideae, vol. 2.
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Johnson, L.A.S. & Wilson, K.L. (1993) Casuarinaceae. Pp. 237-242, in Kubitzki, K., Rohwer, J.G. &
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145-161.
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Li, S.-P. & Hsieh, C.-F. (1997) Seedling morphology of some woody species in a subtropical rain
forest on southern Taiwan. Taiwania 42: 207-238.
Maggia, L. & Bousquet, J. (1994) Molecular phylogeny of the actinorhizal Hamamelidae and
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Martin, P.G. & Dowd, J.M. (1993) Partial sequences of ribosomal RNA of Papilionaceae and related
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Nicolle, D. & Conran, J.G. (1999) Variation in the Eucalyptus flocktoniae complex (Myrtaceae) and
the description of four new taxa from southern Australia. Austral. Syst. Bot. 12: 207-239.
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Prider, J. (1998) The population structure and reproductive biology of Gymnostoma australianum
L. Johnson. Unpublished B.Sc. (Honours) thesis. University of Adelaide.
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Gymnostoma. Pp. 137-147, in Douglas, J.G. & Christophel, D.C. (eds). Proceedings of the 3rd
International Organisation of Palacobolanisls Conference. (A-Z Printers: Melbourne).
Scriven, L.J. & Hill, R.S. (1995) Macrofossil Casuarinaceae: their identification and the oldest
macrofossil record, Gymnostoma antiquum sp. nov. from the Late Paleocene of New South Wales.
Austral. Syst. Bot. 8:1035-1053.
Torrey, J.G. (1983) Root development and root nodulation in Casuarina. Pp. 180-192, in Midgley,
S.J., Turnbull, J.W. & Johnston, R.D. (eds), Casuarina Ecology, Management and Utilization.
(CSIRO: Melbourne).
Torrey, J.G. & Berg, R.H. (1988) Some morphological features for generic characterisation among
the Casuarinaceae. Amer. ]. Bot. 75: 864-874.
Turnbull, J.W. & Martensz, P.N. (1982) Aspects of seed collection, storage and germination in
Casuarinaceae. Austral. Forest Res. 12: 281-294.
Weberling, F. & Leenhouts, P.W. (1966) Systematisch-morphologische Studien an
Terebinthales-Familien (Burseraceae, Simaroubaceae, Meliaceae, Anacardiaceae, Sapindaceae).
Akad. Wiss. Abh., Math.-Natunoiss. Kl. 10: 536-542, 570-571.
Williams, R.F. & Metcalf, R.A. (1985) The genesis of form in Casuarinaceae. Austral. J. Bot. 33:
563-578.
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Manuscript recieved 16 September 1999
Manuscript accepted 22 March 2000
441
The taxonomy of Xanthosia huegelii and closely
related species (Apiaceae: Hydrocotyloideae)
J.M. Hart
Abstract
J.M. Hart (John Ray Herbarium, School of Biological Sciences, Macleay Building M2, The University of
Sydney, NSW 2006, Australia; e-mail: jmh@bio.usyd.edu.au). 2000. The taxonomy o/Xanthosia huegelii
and closely related species (Apiaceae: Hydrocotyloideae). Telopea 8(4): 441-453. Recognition of the three
species Xanthosia huegelii, X. pusilla and X. tasmanica has been inconsistent across their distribution
in southern Australia. As part of a revision of the genus, the history of this group of species is
reviewed and the position of the competing names clarified. Examination of the morphology of
the species, both from living and herbarium material, resulted in only two taxa being recognised,
X. huegelii and X. tasmanica. Both species are more widely distributed than previously thought, due
mainly to the misapplication of the name X. pusilla.
Introduction
Xanthosia huegelii (Benth.) Steud., X. pusilla Bunge and X. tasmanica Domin are small
herbaceous species, commonly occurring in eucalypt forest and heath communities in
southern Australia. These species are distinguished from all other members of the
genus by their trifoliolate leaves with entire segments.
In the course of a revision of the genus, the delimitation of the currently recognised
species was found to be inconsistent. There has been much confusion in distinguishing
the three species from one another, particularly in the application of the competing
names X. tasmanica and X. pusilla, considered as distinct taxa in some treatments
(e.g. Eichler 1986) and a single species, X. pusilla, in others (e.g. Curtis 1963).
This paper aims to clarify the application of the competing names based on field
observations and herbarium material, including examination of the relevant type
specimens.
Taxonomic history
Leucolaena huegelii was described by Bentham in 1837 from specimens collected in the
Swan River and King Georges Sound areas in Western Australia. Bentham described
the species as having trifoliolate leaves with linear segments, and pedunculate
compound umbels with 3-5 rays and two flowers in each umbellule. Steudel (1841:
790-791) transferred the species from the illegitimate genus Leucolaena R. Br. to
Xanthosia Rudge.
Bunge (1845) described two new species, X. pusilla and X. glabrata. Xanthosia pusilla
was described as possessing oblong leaf segments and a sessile, simple umbel with 1-2
flowers. Xanthosia glabrata was described as a glabrous herb with oblong-linear leaf
segments and pedunculate simple umbels with 1-3 flowers. Both species were based
on specimens from Western Australia.
442
Telopea 8(4): 2000
Turczaninov (1849) described X. villosa, a herb with trifoliolate leaves with linear-
cuneate segments and pedunculate, simple umbels of two flowers. However, his type
specimen, Drummond's 4th collection no. 133, was also cited within the same
publication as the type for Dimetopia grandis, the base name for Trachymene grandis
(Turcz.) Rye (Rye 1999). Examination of photographs of this collection confirms it to be
a Trachymene having simple umbels with numerous flowers. A survey of Turczaninov's
type specimens at KW found the intended type of X. villosa to be Drummond's 4th
collection no. 139 (Marchant 1990), a specimen collected at Swan River in Western
Australia which conforms to Turczaninov's description. As it can only be considered
illogical for Turczaninov to have intended to publish these apparently homotypj c
synonyms in the same work, the type citation for X. villosa should be treated as a
typographical error. Thus the type for the name X. villosa is here lectotypified as
Drummond's 4th collection no. 139.
Bentham (1867) listed X. villosa in synonymy under X. pusilla, and X. glabrata was
reduced to varietal status as X. pusilla var. glabrata. The two species he recognised in
his treatment, X. huegelii and X. pusilla, were separable by the inflorescence being a
simple umbel (in X. pusilla) or a compound umbel (X. huegelii). Xanthosia huegelii was
considered to be restricted to Western Australia, whereas X. pusilla was stated to occur
in Western Australia, South Australia, Victoria and Tasmania.
Xanthosia tasmanica was based upon specimens collected at Circular Head and
Georgetown in Tasmania (Domin 1907). This species was described as having-
elliptical leaf segments with the lateral segments sometimes bifid; subsessile, 1-2-
flowered umbels; and five-ribbed fruit. Ostenfeld (1921) reported X. tasmanica as
occurring in Western Australia and agreed that the species was different from
X. pusilla. However, X. tasmanica has not ever been subsequently considered to occur
in Western Australia.
Recently Keighery informally divided the X. huegelii specimens in PERTH (sheet
annotations) into three unpublished subspecies; X. huegelii 'subsp. huegelii' with linear
leaflets and a compound umbel; X. huegelii 'subsp. aurea', a variant occurring in the
Eneabba to Gingin area north of Perth, having inflorescences with more numerous
flowers and broader bracteoles; and an unnamed subspecies 'Southern' with broadly
elliptic leaflets, occurring in coastal areas in the south of the state. y
Discussion
The main features used by previous workers to distinguish the three taxa (X. huegelii
X. pusilla and X. tasmanica) were leaf segment shape and inflorescence elaboration
Specimens could be divided into two groups based on leaf segment shape; those with
elliptic leaf segments, and those with linear, oblong, lanceolate or oblanceolate leaf
segments (Figs 1 and 2). Within the group with elliptic leaf segments (Fig. 1), the lateral
segments of the leaves on young plants are often partly to fully divided into two lobes
he leaves of this group are covered by a sparse covering of long multicellular
multisenate trichomes. The group with linear, oblong, lanceolate or oblanceolate leaf
egments (Fig 2) is densely villous to glabrous, sometimes becoming glabrous with
age. Very rarely, the segments may be broadly toothed (Fig. 2g). The stems and taproot
of many o the plants with this leaf form excorticate with a£ As t^ToesnoZcul
ev,VW 8 r P u K ° rthC neW gr ° Wth fr ° m Whkh the infl orescences arise, it is not always
H her ?T m S P ecimens - The two leaf forms occur at the same locality on
separate individuals without the presence of intermediate leaf forms ( Hart 95108 &
Hart, Taxonomy of Xanthosia huegelii
443
Fig. 1. Leaf silhouettes of Xanthosia tasmanica showing variation within and between individuals:
a, basal; b, mid-branch; c, subtending inflorescence (from Hart 95109); d, basipetal, e, acropetal
(from Hart 369). All leaves actual size.
Fig. 2. Leaf silhouettes of Xanthosia huegelii showing variation within and between individuals,
a, basipetal; b, acropetal (from Hart 95055); c, basipetal; d, acropetal (from Hart 381);
e, basipetal; f, acropetal (from Hart 95134); g, basipetal; h, acropetal (from Hart 95142). All leaves
actual size.
The inflorescences vary in the extent of elaboration (simple or compound) and size
(rays up to 5 mm long)"in individuals of both leaf forms. Within the group with elliptic
leaf segments, inflorescences vary from one- to four-rayed with one flower per ray
(umbellule) and with one flower sometimes present between the rays. The flowers are
all bisexual and the involucral bracts and bracteoles are all linear. The group with
linear, oblong, lanceolate or oblanceolate leaf segments also has 1-4 rays in the
inflorescences with 1 or 2 bisexual flowers in each umbellule, sometimes with one
flower between the rays. At the north-western extreme of its distribution, in the
Eneabba area north of Perth, the inflorescences are larger with broader, elliptic
bracteoles, and up to six bisexual and/or male flowers in each rayed umbellule and up
to three flowers between the rays. Continuous variation in flower number and
between elliptic and linear involucral bracts and bracteoles occurs such that division
of this group is largely arbitrary. The flowers of the two leaf forms are morphologically
consistent, while the fruit vary in rib number; one form has 5-7 ribs, whereas the other
has 5-9 ribs.
444
Telopea 8(4): 2000
The two leaf forms are here considered to be sufficiently distinct to be treated as two
separate — but closely related — species, which accord with the types of X. tcismanica
(with elliptic leaf segments), and X. huegelii (with linear, oblong, lanceolate or
oblanceolate leaf segments).
Keighery (1982) determined the chromosome numbers for many Western Australian
Apiaceae, and lists all but one species of Xanthosia as having n=10. Only X. pustlla was
found to have n=20. The chromosome voucher specimens were redetermined in line
with the two species recognised here. The chromosome numbers for X. huegelii are
both «=10 (seven specimens) and »=20 (10 specimens), and for X. tcismanica are also
n=10 and ii=20 (one specimen and two specimens respectively; Table 1).
Table 1. Current determinations of chromosome voucher specimens following the author's
concepts.
Chromosome numbers taken from Keighery (1982) and from sheet information where available.
Locality
Xanthosia huegelii
n
Voucher and Herbarium
16 km NW of Cataby, W.A.
10
G.J. Keighery s.n., PERTH 3620700
170 km N of Perth on Brand Highway, W.A.
10
G.J. Keighery 97, PERTH 4878728
5 km S of Corbanup River Store,
Busselton/Margaret River Road, W.A.
10
G.J. Keighery 690, KPBG
8.5 miles [13.6 km] N of Eneabba, W.A.
10
G.J. Keighery 1338, PERTH 3620506
Cannington Swamp, W.A.
10
G.J. Keighery 2102, PERTH 3620581
24 miles [38 km] S of Nannup on Pemberton
road, W.A.
10
G.J. Keighery 2166, PERTH 3620891
1.2 miles [1.9 km] W of Brand Highway
on Mimegarra road, W.A.
10
G.J. Keighery 2753, PERTH 3620530
Inlet River, Walpole to Manjimup, W.A.
20
G.J. Keighery 351, PERTH 3621278
32 km N of Walpole, W.A.
20
G.J. Keighery 81, PERTH 4878744
Mt Burnett Wittenoom Hills
c.70 km NE Esperance, W.A.
20
G.J. Keighery 424, PERTH 3621286,
CANB 370637
Mt Merivale c. 30 km E of Esperance, W.A.
20
G.J. Keighery 431, PERTH 3621421
395 mile peg, Ravensthorpe to Esperance road, W.A.
20
G.J. Keighery 492, PERTH 1867547
47.2 miles [75.5 km] E of Albany
on Jerramungup road, W.A.
20
G.J. Keighery 494, PERTH 3621243
22 miles [35 km] E of Esperance, W.A.
20
G.J. Keighery 653, KPBG
10.2 km N Walpole on Manjimup road, W.A.
20
G.J. Keighery 677, PERTH 3621189
Nicholls road Mt Lofty Ranges, S.A.
20
G.J. Keighery 2248, PERTH 3621383
Mt Merivale E of Esperance, W.A.
Xanthosia tasmanica
20
G.J. Keighery 2887, PERTH 3621448
18 miles [29 km] S of turnoff to Busselton
on Yallingup to Karridale Road, W.A.
10
G.J. Keighery 2165, PERTH 3620859
46 km W of Ravensthorpe, W.A.
20
G.J. Keighery 104, PERTH 4878752
25 km NW Denmark on Walpole Road, W.A.
Not located: listed as X. pusilla by Keighery (1982)
20
G.J. Keighery 673, PERTH 3628590
Nannup, W.A.
20
G.J. Keighery s.n.
24 km S of Manjimup, W.A.
20
G.J. Keighery 78
Hart, Taxonomy of Xanthosia huegelii
445
The specimens which were found to be n=10 occur in the south-west of Western
Australia, north from Nannup, whereas specimens from the south coast of Western
Australia, east from Nannup, and the one specimen from South Australia are n= 20
(Fig. 3). However, no consistent morphological differences were found to correlate
with the different chromosome numbers in either species.
Xanthosia huegelii was previously considered to be restricted to Western Australia;
however as here circumscribed it is also found in South Australia and Victoria, where
it was previously treated as X. pusilla. Xanthosia tasmanica also occurs in Western
Australia, where it constitutes Keighery's unnamed subspecies 'Southern' within
X. huegelii, as well as occurring in South Australia, Victoria, New South Wales and
Tasmania. Table 2 summarises the names previously applied to each species in each
state of their distribution.
Table 2. Summary of previously applied names to X. huegelii and X. tasmanica in each State of
their distribution.
State
Xanthosia huegelii
Xanthosia tasmanica
Western Australia 1
X. huegelii subsp. huegelii
X. huegelii subsp. aurea MS.
X. pusilla
X. huegelii subsp. 'Southern' MS.
South Australia 2
X. pusilla
X. tasmanica
Victoria 3
X. pusilla
X. pusilla
Tasmania 4
-
X. pusilla
New South Wales 5
_
misidentified as X. pilosa
1 Wheeler (1987) and X. huegelii subspecies manuscript names as sorted in PERTH,
2 Eichler (1986), 3 Willis (1978), 4 Curtis (1963), 5 Brooks and Powell (1992).
Key to the trifoliolate-leaved herbaceous species of Xanthosia
1 Stems excorticating when aged; leaf segments linear to oblong or lanceolate; fruit with
5-9 ribs . 1. X. huegelii
1* Stems not excorticating when aged; leaf segments elliptic, the lateral segments
sometimes bifid; fruit with 5-7 ribs . 2. X. tasmanica
Taxonomy
1. Xanthosia huegelii (Benth.) Steud., Nomencl. hot. 2: 791 (1841).
Leucolaena huegelii Benth. in Endl. et al., Enum. pi, 55 (1837).
Type: Western Australia: Swan River, Hiigel (lecto, here chosen, K!; isolecto W, photo!).
Bentham's protologue cites 'King Georges Sound et Swan River. (Hiigel)', however,
only material labelled 'Swan River, Hiigel' has been located. As this material clearly
fits the details of the protologue it is here designated as the lectotype.
Xanthosia glabrata Bunge in Lehm., PI. Preiss. 1: 290 (1845).
Xanthosia pusilla var. glabrata (Bunge) Benth., FI. Austral. 3: 361 (1867).
Type: In umbrosis sylvae ad fluvium Canning, Perth, Western Australia, Nov 1841,
Herb. Preiss. No. 2076 (holo P; iso MEL!).
446
Telopea 8(4): 2000
Fig 3. Distribution of chromosome voucher specimens for X. huegelii (• n=10, O „=20) and
X. tasmamca Domin (■ n=10, □ n=20). 3 ' J> anct
Kilometres
Hart, Taxonomy of Xanthosia huegelii
447
Xanthosia pusilla Bunge in Lehm., PI. Preiss. 1: 291 (1845).
Type: In arenosis humidis umbrosisque ad fluvium King's-River, Plantagenet, Western
Australia, 14 Oct 1840, Herb. Preiss. No. 2078 (holo P; iso MEL!).
Xanthosia villosa Turcz., Bull. Soc. Imp. Nat. Moscou 22(2): 32 (1849).
Type: Western Australia: Swan River, /. Drummond IV. n. 139 (lecto, here chosen, KW,
photo!; isolecto BM!, K!). Turczaninov's type designation is discussed under
Taxonomic History.
Erect or ascending perennial herb, 5-20 cm high. Plant near glabrous to densely
villous, the stem excorticating when aged. Leaves cauline, trifoliolate. Petiole
sheathing shortly at the base to entirely sheathing, 1-7 mm long. Leaflets linear,
oblong, lanceolate or oblanceolate, the segments equal, very rarely lobed, 4-38 mm
long, 1-10 mm wide; margins entire; apex acute. Inflorescence a compound umbel with
1-4 rays, sometimes contracted and appearing simple, 1-6 flowers per ray and up to 3
flowers between the rays; the flowers all bisexual to bisexual and male. Rays terete, to
5 mm long. Involucral bracts 1-4, elliptic or linear, foliaceous, green, longer than the
rays, 3-9 mm long, 1-4 mm wide; apex acute or obtuse. Bracteoles 2 or 3, elliptic or
linear, foliaceous, green, shorter to longer than the flowers, 4-6 mm long, 1-3 mm
wide; apex acute or obtuse. Inflorescences sessile or pedunculate; peduncles to 25 mm
long. Flowers pedicellate. Sepals ovate, 1.1-2.9 mm long, 0.7-1.4 mm wide, green; base
truncate; apex acute, glabrous. Petals equal to sepals, 1-1.2 mm long, 0.3-0.6 mm wide,
white; midrib adaxially keeled and bridged; apex acute or obtuse; appendage smooth.
Stamens approximately equal to the perianth. Nectaries raised and prominent, 0.4-0.7 mm
high, cream, hirsute. Styles 0.7-1.9 mm long. Ovary hirsute. Male flowers different
from bisexual ones in having undeveloped inconspicuous ovary, with styles barely
protruding above nectaries. Fruit brown, ovoid, 2.3—2.8 mm long, 1.9—2.3 mm wide,
0.4-0.7 mm deep. Monocarps villous on the summit, rarely glabrous, oblong in
transverse section, 5-9 ribbed; the ribs keeled. Chromosome number >7=10 and n=20
(Keighery 1982; Table 1). (Fig. 4).
Flowering: most of the year.
Notes: Xanthosia huegelii is a variable species particularly in indumentum and
inflorescence size. A variant with dense greyish indumentum, broader bracteoles and
a greater number of flowers per umbellule occurs in Western Australia around
Eneabba and south to Perth, with intergrading forms south of Perth. Placement of
specimens into any form-groups is largely arbitrary.
Distribution: coast and adjacent ranges in Western Australia (between Eneabba and
Esperance), South Australia (on the Eyre Peninsula, Kangaroo Island and east from the
Lofty Ranges) and south-western Victoria (the Grampians to Wilsons Promontory)
(Fig. 5).
Ecology: widespread in forest and heath usually on sandy soils.
Selected Specimens (over 200 examined): Western Australia: 6 miles [c. 10 km] N of junction of
Eneabba South road with Coorow-Green Head road. Chapman s.n., 30 Aug 1977 (PERTH 3260611);
0.5 km N of Marchagee Track W of Brand Highway, Cranfield 8892 & Kabay, 3 Sept 1993 (PERTH);
14.6 km along the Mooliabeenee road from the Great Northern Highway towards Gingin, Hart
95055, 25 Oct 1995 (PERTH, SYD); Delta road, Walpole-Nornalup National Park, Hart 95108, 8 Nov
1995 (PERTH, SYD); Porongorups, 1.3 km along Angwin Park road from Bolganup road. Hart
95129, 13 Nov 1995 (PERTH, SYD); 2.8 km along the South Coast Highway from Chester Pass road
towards Jerramungup, Hart 95134, 15 Nov 1995 (SYD); Lucky Bay, Cape Le Grand National Park,
Hart 95142, 18 Nov 1995 (SYD); Eneabba, Humphreys 342, 21 Sept 1966 (PERTH); Keighery 5366,
Oct 1975 (PERTH); Gold Holes, Chester Pass, Stirling Ranges, Keighery 5533, 19 Oct 1982 (PERTH);
Goode Beach area, Torndirrup National Park, 18 km S of Albany, Keighery 8513,
5 Nov 1986 (PERTH); Cape Le Grand National Park E of Esperance, Royce 8713, 21 Oct 1969
448
Telopea 8(4): 2000
Fig. 4. Xanthosia huegelii. a, branchlet; b, c, petals, (b, adaxial view; c, side view); d, inflorescence
1 ScITh rayS ^ lth T fl ° Wer betWeen the ra y s; e ' leaf ; { ' bisexual flower; g, fruit (from
Hart 379). Scale bar: a = 25 mm; b, c = 0.8 mm; d = 4 mm; e = 20 mm; f = 1.2 mm; g =1.5 mm.
Hart, Taxonomy of Xanthosia huegelii
449
(PERTH); Blue Waters, Collie, Saffrey 1763, 1 Nov 1979 (CANB, K, PERTH); Helena Valley, Seabrook
271, 25 Sept 1977 (PERTH); Station road Walpole-Nornalup National Park, Wheeler 3159, 13 Aug
1992 (PERTH).
South Australia: Hundred of Wanilla, Section 124 Reserve [c. 25 km NNW of Port Lincoln], Alcock
798, 17 Oct 1965 (AD); Big Heath National Park c. 1.5 km S and parallel to the fire break, mid¬
narrow neck (c. 25 km SW Naracoorte), Alcock 2913, 4 Nov 1969 (AD); 4 km SW of Bradbury on
the western side of Mt Bold road. Dormer 9721, 3 Nov 1983 (AD); Kangaroo Island on Bark Hut
road near Amen Comer, Jackson 3167, 21 Nov 1993 (AD); Start of track 6.6 km S of Ashbourne by
road, Coxs Scrub Conservation Park, Hart 381, 23 Nov 1996 (AD, SYD).
Victoria: 6.5 km N of Portland-Nelson road on Wrights Swamp road, Hart 379, 21 Nov 1996 (MEL,
SYD); Slopes between Great Ocean road and Harvey Street Anglesea, Smith 59/288, 1 Nov 1959 (MEL).
Fig. 5. Distribution of Xanthosia huegelii (•) based on locality information from herbarium specimens.
450
Telopea 8(4): 2000
2. Xanthosia tasmanica Domin, Repert. Spec. Nov. Regni Veg. 4: 298 (1907) non Gand.,
Bull. Soc. Bot. France 65: 32 (1918).
Type. Circular Head, Tasmania, R. Gunn 1837 / 879 (lecto, here chosen, K!, the largest
specimen in the top left corner). Domin cited two specimens in the protologue for
X. tasmanica ; ^'prope George Town (Ig. Gunn 1842 sub no. 1122) et Circular Head
(Ig. Gunn 183/ sub no. 879) (H. Kewense)'. This material, comprising nine elements, is
mounted on a single sheet at K. Thus there is some ambiguity as to which collection(s)
were from George Town and which were from Circular Head. The largest element, in
the top left corner, has a determinavit label below it by Dr Hansjoerg Eichler (dated
1961) indicating his intention to lectotypify this plant (which he never published).
The nearest locality label to this plant is for the Circular Head collection. This largest
p ant is designated as lectotype (in accordance with Dr Eichler's intentions) as it is the
best specimen on the sheet.
Erect or ascending perennial herb to 20 cm high. Plants covered by sparse, long,
spreading hairs. Leaves cauline, trifoliolate. Petiole sheathing shortly at the base,
sheaths 1-8 mm long. Petiole to 30 mm long, angular, grooved above or flattened.
Leaflets elliptic, the segments equal or the lateral leaflets split or parted almost to the
base, 4-15 mm long, 3-8 mm wide, shortly petiolulate; margins entire; apex acute.
iif orescence a compound umbel with 1-4 rays, sometimes contracted and appearing
simple, one flower per ray and up to one flower between the rays; the flowers all
bisexual Rays terete, 1-3 mm long. Involucral bracts 1-4, linear, foliaceous, green,
onger than the rays, 3-4 mm long, 1 mm wide; apex acute. Bracteoles two or three,
mear, foliaceous, green, shorter than flowers, 3-1 mm long, 1 mm wide; apex acute.
Inflorescences sessile or pedunculate; peduncles to 5 mm long. Flowers pedicellate.
Sepals ovate, 2-2.5 mm long, c. 1 mm wide, green; base truncate; apex acute, glabrous.
etals shorter than sepals, c. 1.2 mm long, c. 0.5 mm wide, white; midrib adaxially
keeled or keeled and bridged; apex acute; appendage smooth. Stamens shorter than
§f, ,ant , h ^ eCtan f S raiS6d and P rominent ' c. 0.3 mm high, white or yellow, hirsute.
•jn } 5 ong ‘ ° var y hirsute. Fruit brown, ovoid, c. 2.1 mm long, c. 2.3 mm
wide, 06 mm deep. Monocarps hirsute on the summit, elliptic in transverse section,
lie - r ! h^i'QooV'u? ^w? d ° r n0t elevated - Chromosome number n =10 or > 2=20
(Keighery 1982; Table 1). (Fig. 6).
Flowering: most of the year, peaking in spring.
Distribution: found in south-west Western Australia, Fleurieu Peninsula and
angaroo Island in South Australia, the Grampians and coastal Victoria to the far
south coast of New South Wales, and coastal Tasmania (Fig. 7).
Ecology: occurring mainly in coastal areas in heath or forest on sandy soils.
Seated Specimens (60 examined): Western Australia: Delta road, Walpole-Nornalup National
673 12 St 1974,f° ); 25kmNW Denmark Walpole road [„=20], Keighery
K ffioS Tto S™i “ lo Bus “ l,on on Y! "" ns " p ,o
Xnv M UStr ? i ia: f H 6 n km NW Pardana ' Kangaroo Island, Lang & Maguire NPKI30860, 19 Nov 1989
(AD), Mount Lofty Range, Spring Mount c. 8 km SE of Myponga, Whibley 1710 ,23 Nov 1966 (AD).
^*^^1991 ^AD^Hn 31 ^b rwff/i' °'° n 8 National P ‘ lrk near mouth of Seal Creek, Albrecht 4846,
1996 (MEL, SYD) °' M ^ MarIo ' Ca P e Conran road at Point Ricardo turnoff. Hart 369, 18 Nov
(kXmEGNSW; Sc)" 1 CaPe Lighth0USe 26 km SE of Eden ' Cov ™y 5811 & Armstrong, 16 Oct 1974
Mm **• 22 1980 <HO 39283,: Healh —• «*<*«.
Hart, Taxonomy of Xanthosia huegelii
451
Fig. 6. Xanthosia tasmanica: a, branchlet; b, inflorescence; c, d, leaves showing variation;
e, f, petals, (e, adaxial view; f, side view); g, bisexual flower; h, fruit (from Hart 95109).
Scale bar: a = 40 mm; b = 5 mm; c, d = 20 mm; e, f = 0.75 mm; g = 1.6 mm; h = 1.5 mm.
452
Telopea 8(4): 2000
Fig. 7. Distribution of Xanthosia tasmaiiica (•) based on locality information from herbarium specimens.
Kilometres
Hart, Taxonomy of Xanthosia huegelii
453
Acknowledgments
The directors and staff of AD, CANB, CBG, HO, KPBG, MEL, NSW and PERTH for
access to and/or loans of specimens, particularly Neville Marchant for allowing me to
examine his photos of the Xanthosia villosa type in KW. Murray Henwood and Bob
Makinson are thanked for critical review of the manuscript.
References
Bentham, G. (1837) Umbelliferae. Pp. 55-56 in Endlicher, S.L., Fenzl, E., Bentham, G. and Schott,
H.W. (eds), Enumeratio plantation qnas in Novae Hollandiae ora austro-occidentali ad fluvium
cygnorum et in sinu Regis Ceorgii collegit Carolus Liber Baron de Hiigel. (Fr. Beck: Vienna).
Bentham, G. (1867) Flora Australiensis, vol. 3. (L. Reeve: London).
Brooks, A.K. & Powell, J.M. (1992) Xanthosia (Apiaceae). Pp. 94-96 in Harden, G.J., (ed.) Flora of New
South Wales, vol. 3. (New South Wales University Press: Sydney).
Bunge, A.A. von (1845) Umbelliferae. Pp 282-295 in Lehmann, J.G.C., Plantae Preissianae, vol. 1.
(Meissner: Hamburg).
Curtis, W.M. (1963) The Student's Flora of Tasmania, Part 2. (Govt Printer: Tasmania).
Domin, K. (1907) Umbelliferae novae extraeuropaeae. 1. Repert. Spec. Nov. Regni Veg. 4: 298.
Eichler, H. (1986) Umbelliferae (Apiaceae). Pp. 979-1010 in Jessop, J.P. & Toelken, H.R. (eds), Flora
of South Australia. Part II Leguminosae — Rubiaceae. (South Australian Government Printing
Division: Adelaide).
Keighery, G.J. (1982) Chromosome numbers of Western Australian Apiaceae. J. Roy. Soc. Western
Australia 65:143-145.
Marchant, N.G. (1990) The contribution of the Russian botanist Turczaninov to Australian plant
taxonomy. Pp. 121-130 in Short, P.S. (ed.). History of Systematic Botany in Australia. (Australian
Systematic Botany Society Inc.: Melbourne).
Ostenfeld, C.H. (1921) Contributions to West Australian Botany. Part III. Additions and notes to
the Flora of extra-tropical W. Australia. Biol. Meddel. Kongel. Danske Vidensk. Selsk. 3(2): 101,102.
Rye, B.L. (1999) A taxonomic revision of the many-flowered species of Trachymene (Apiaceae) in
Western Australia. Nuytsia 13:193-232.
Steudel, E.G. von (1841) Nomenclalor Botanicus, edn 2,2.
Turczaninov, N.S. (1849) Bull. Soc. Imp. Nat. Moscou 22: 3-38.
Wheeler, J.R. (1987) Apiaceae. Pp. 501-518 in Marchant, N.G., Wheeler, J.R., Rye, B.L., Bennett, E.M.,
Lander, N.S., & Macfarlane, T.D. (eds), Flora of the Perth Region, Part 1. (Western Australian
Herbarium, Department of Agriculture: Perth).
Willis, J.H. (1978) A Handbook to Plants in Victoria, vol. 2. (Melbourne University Press: Melbourne).
Manuscript received 22 April 1999
Manuscript accepted 22 March 2000
455
Actinotus periculosus (Apiaceae): a new
perennial species from eastern Australia
M.J. Henwood
Abstract
M.j. Henwood (John Ray Herbarium, School of Biological Sciences (M2), The University of Sydney, NSW
2006, Australia). 2000. Actinotus periculosus (Apiaceae): a new perennial species from eastern Australia.
Telopea 8(4): 455-459. A new perennial species, Actinotus periculosus, is described from the
Leichhardt District of Queensland. A. periculosus is allied to A. helianthi (New South Wales and
Queensland) and A. schumrzii (Northern Territory), but is separable from these species by its larger,
deeply dissected leaves and relatively long gynoecial column. A key to all species of Actinotus from
eastern and central Australia is provided.
Introduction
Actinotus as it is currently circumscribed comprises 17 species. The genus is most
diverse within Australia, and is represented by a single endemic species, A. novae-
zelandiae (Petrie) Petrie, in New Zealand. Within the Australian Apiaceae, Actinotus is
recognisable on account of its compressed monocarpic fruit, and its often showy
involucral bracts forming pseudanthial umbels or capitula. The single-seeded
monocarp is apparently derived from the more common bicarpellate gynoecium, for
it initially contains two ovules, one of which subsequently fails to develop beyond a
rudimentary structure (Magin 1977).
One of the most widespread Australian perennial species is A. helianthi, which ranges
from Merimbula (36°54'S, 149°54'E) in southern New South Wales to Rolleston
(24°27'S, 148°37'E) in Queensland. Two leaf forms can be discerned within
A. helianthi. Plants from coastal heaths (at the top of sea cliffs or on stable coastal
dunes) are more robust and have less dissected leaves than plants from woodland
habitats. When grown in common gardens these morphological differences are
maintained, thereby appearing to be genetically fixed (von Richter pers. comm.).
Together with A. schwarzii (restricted to the cliff faces of the MacDonell Ranges;
Northern Territory), Actinotus helianthi constitutes a morphologically distinct group
within the genus. Both species are morphologically closely allied and share several
unique, apparently derived characters namely: more or less filamentous petals, an
elongated restriction (column) above the ovary and long horn-like nectaries inserted
at the base of the styles.
Actinotus helianthi reaches its northern-most limit in the vicinity of the Carnarvon
range (Queensland). The Carnarvon range is formed in large part by the outcropping
of Jurassic, Precipice sandstone which marks the northern boundary of the Surat basin.
The horizontally bedded Precipice sandstone is deeply dissected and forms vertical
cliffs up to 200 m high. A perennial taxon closely resembling the 'woodland' form of
A. helianthi has been recorded from the cliffs of the Carnarvon range and is here
described as a new species.
456
Telopea 8(4): 200q
Taxonomy
Actinotus periculosus Henwood, sp. nov.
Actinotus helianthi Labill. et A. schwarzii F. Muell affinis. A. helianthi foliis magis penitu s
et subtiliter dissectis, collis longioribus differt. A. schwarzii foliis majoribus stipellis
basalibus destitutus, collis brev'ioribus, petalis filamentosis differt.
Type: Queensland: Leichhardt District: 15 km N of 'The Tombs' on Marlong Creek,
Mt Moffat region of Carnarvon National Park, M.J. Henwood 511, 19 Oct 1998 (holo
SYD; iso BRI, NSW).
Plants andromonoecious, erect to ascending, perennial, to 20-50 cm high. Stems
branching, covered in dense white felted indumentum. Leaves cauline, discolorous,
alternate, temately dissected, petiolate. Sheaths 6-9 mm long, sericeous and felted.
Petioles 15-54 mm long, adaxially grooved. Laminas 20-100 mm long, 15-70 mm
wide, finely dissected. Primary segments elliptic to linear, (20-)30-65 long, 1—5(—6) mm
wide, apices acute or obtuse. Umbels capitulum-like, pseudanthial, borne in
monochasial cymes, subtended by trifid bracts. Peduncles 30-90 mm long. Umbels
15-30 mm in diameter, 94-310 flowered. Involucral bracts 9-12, apparently arranged
in a single whorl; 25-55 mm long, 5-16 mm wide; lamina hairy, white with green tips,
venation distinct. Bisexual flowers borne centrally in 'capitula' and surrounded by a
peripheral ring (or rings) of male flowers. Anthesis proceeds centrifugally in each
umbel. Bisexual flowers protandrous. Pedicels 4-4.5 mm long, articulated at junction
with the gynophore, sparsely villous. Gynophores 3.4-4 mm long, villous. Sepals
conspicuous, petaloid, ovate, 1—1.5 mm long, 0.5-0.75 mm wide, white to cream, bases
truncate, apices acute, villous. Petals inconspicuous, filamentous, 0.8-0.95 mm long,
glabrous. Stamens 5; filaments 1.7-2.5 mm long; anthers dorsifixed, 0.4-0.5 mm long'
Stylopodia 0.5-0.9 mm high, glabrous. Nectaries 2, each a horn-like projection inserted
directly below each style, 0.3 mm long, dark purple, glabrous. Ovaries unilocular,
villous, surmounted by a column; column 2.4-3 mm long, villous. Styles 2, arranged'
parallel to the plane of ovary compression, 1.4-2 mm long. Male flowers characterised
by a much reduced (non-functional) gynoecium, otherwise similar to bisexual flowers
Mature fruit not seen. (Fig. 1).
Distribution and habitat: endemic to the Leichhardt District of Queensland
Restricted to sandstone cliff faces (Fig. 2).
Notes: The characteristically deeply dissected leaves with narrow, linear segments of
A. periculosus do not correspond to either the woodland or the heath leaf forms of
A. helianthi. Individuals of A. periculosus are further separable from A. helianthi by the
former's longer gynoecial columns (2.4-3.0 mm long). Based on observations from one
flowering season, A. helianthi and A. periculosus appear to be reproductively isolated
when in sympatry — for example at Spyglass Mountain, in the Salvator Rosa region
ot Carnarvon National. In such localities A. helianthi is fruiting whilst A. periculosus is
at early anthesis.
Actinotus periculosus is likewise easily separable from A. schwarzii. Like A. helianthi
leaves of A periculosus lack the pair of basal basiscopic, stipel-like lobes that
2t ra lT S , e ! he l f aves J oM - schwarzii. In addition, the length of the gynoecial columns
ot both A helianthi and A. periculosus are not as long as those of A. schwarzii (> 3 0 mm
m Sot Pet f °! A schwarzii are a P icall y broader than those of the other species
1 owell (1992) and others have erroneously described A. helianthi as apetalous). Given
e apparent reproductive isolation of A. periculosus from A. helianthi when in
sympatry, combined with obvious differences in reproductive and vegetative
A°heUanth P *“ ^ A periculosus does ' in fact ' warrant formal segregation from
Henwood, Actinotus periculosus sp. nov.
457
Flowering: October and November
Etymology: from the Latin periculosus, dangerous, in reference to the difficulty
experienced in obtaining representative specimens of this species from sandstone cliffs.
Conservation status: this species is not regarded as rare and is adequately protected.
Selected specimens: Queensland: Leichhardt: 'The Tombs', c. 143 km from Injune, Mt Moffatt
section of Carnarvon National Park, Henwood 508,18 Oct 1998 (BR1, NSW, SYD); 15 km N of 'The
Tombs' on Marlong creek, Mt Moffat region of Carnarvon National Park, Henwood 511 ,19 Oct 1998
(BR1, NSW, SYD); Lookout at Lonesome National Park, Henwood 513,19 Oct 1998 (BRI, NSW, SYD);
'Balloon Cave', Carnarvon Gorge, Carnarvon National Park, Henwood 517 ,21 Oct 1998 (BRI, NSW,
SYD); 'Spyglass Peak' Salvator Rosa section of Carnarvon National Park, Henwood 518 ,21 Oct 1998
(BRI, NSW, SYD); c. 100 m north of Get Down track, Robinson National Park, Henwood 520 ,23 Oct
1998 (BRI, NSW, SYD).
J
Fig. 1, Actinotus periculosus (from Hemoood 511). a, foliose upper portion of branch with
inflorescence; b, hermaphrodite flower; c, apex of gynoecial column with 3 sepals, 4 petals and
2 stamens removed (trichomes not illustrated). Scale bar: a = 100 mm; b = 5 mm; c = 4 mm.
458
Telopea 8(4): 2000
Key to eastern and central Australian species of Actinotus
1 Leaves dissected; flowers in capituloid umbels; bracts > 19 mm long; gynophores
present; columns present; petals filamentous . 2
2 Leaf laminas without basal, basiscopic lobes; column < 3 mm long; petals hair-like,
not apically expanded ..... 3
3 Leaf segments long-linear; column 2.4-3 mm long. Qld. A. periculosus
3* Leaf segements broadly linear; column < 1 mm long. Vic., N.S.W., Qld.
. A. helianthi
2* Leaf laminas with basal, basiscopic lobes; column > 3 mm long; petals slightly
apically expanded. N.T. . A. schwarzii
1* Leaves entire, toothed or dissected; flowers in capitula or capituloid umbels;
bracts < 10 mm long; gynophores absent; columns absent; petals, when present,
spathulate . 4
4 Plants cushion-forming to 7 cm high; capitula with 5-12 flowers, not pseudanthial;
bracts 1.5-3.0 mm long; flowers sessile; petals absent; fruit weakly laterally
flattened....... g
5 Leaves entire, crenulate or shallowly trilobed; sepals free. 6
6 Leaves entire; stamens 2-3. Tas. A. suffocatus
6 * Leaves crenulate or shallowly trilobed; stamens 5. Tas., Vic.
. A. bellidioides
5* Leaves dissected; sepals basally connate; stamens 2. Tas. A. moorei
4* Plants erect, procumbent or decumbent, never cushion-forming, usually > 5 cm
high; umbels or capitula with 16-68 flowers, pseudanthial; bracts 4 . 0-10 mm long;
flowers pedicellate; petals absent or present; fruit strongly laterally flattened
. 7
7 Plants erect to procumbent; sepals free or connate for most of their length;
peduncles 45-150 mm long. g’
8 Involucral bracts 1-whorled; petals absent, sepals free. N.S.W.
... A. minor
8 * Involucral bracts 2-whorled; petals spathulate; sepals connate for most of
their length. Vic., N.S.W. . A. forsythii
7* Plants spreading to weakly erect; sepals basally connate; petals spathulate-
peduncles 1-8 mm long. g
9 Fruit winged, fringed with short trichomes; stamens 3; involucral bracts
basally connate. N.S.W., Qld. A. paddisonii
9* Fruit not winged, fringed with long trichomes; stamens 5; involucral bracts
free. N.S.W., Qld. A . gibbonsii
Henwood, Actinotus periculosus sp. nov.
459
Acknowledgments
The assistance of Gavin Enever and Frank Carter (Queensland National Parks and
Wildlife Service), Jenny Hart and Selga Harrington (SYD), Tony Bean (BRI) and
Maryann Henwood is gratefully acknowledged. Lotte von Richter generously
provided unpublished data on A. helinnthi and permitted access to plants of
A. periculosus and A. helianthi cultivated at Mount Annan (Royal Botanic Gardens
Sydney). This research was funded in part by the Australian Biological Resources
Survey.
References
Magin, N. (1982) Bliitenmorphologische untersuchungen an Actinotus Lab. (Hydrocotyloideae)
unter besonderer beriicksichtigung des gynoeciums. Pp.749-764 in Cauwet-Marc, A.-M. &
Carbonnier, J. (eds), Les Ombelliferes: contributions pluridisciplinaires a la systematique.
[Monographs in Systematic Botany from the Missouri Botanic Garden, vol 6.] (Braun-Brumfield
Inc.: Ann Arbor).
Powell, J.M. (1992) Apiaceae. Pp. 87-116 in Harden, G.J. (ed.). Flora of New South Wales, vol. 3.
(New South Wales University Press: Kensington).
Manuscript received 18 January 2000
Manuscript accepted 10 April 2000
461
The lichen genera Phaeographis and
Phaeographina (Graphidaceae) in Australia
1: Species based on Australian type specimens
Alan W. Archer
Abstract
Archer, Alan W. (Royal Botanic Gardens Sydney, Mrs Macquaries Road, Sydney NSW 2000, Australia )
2000. The lichen genera Phaeographis and Phaeographina ( Graphidaceae) in Australia 1. Species based
on Australian type specimens. Telopea 8(4): 461-475. The types and other specimens of 14 Australian
species in the genus Phaeographis (Graphidaceae) and 3 types and other specimens of Australian
taxa in the genus Phaeographina (Graphidaceae) were examined. Phaeographina caesiopruinosa var.
monospora Miill. Arg. is raised to species level as Phaeographina muelleri A.W. Archer. Phaeographis
subcompulsa Mull. Arg., Phaeographis cinerascens Miill. Arg., Phaeographis inscripta Mull. Arg. and
Graphis aulacothecia C. Knight are taxonomic synonyms of Phaeographis australiensis Mull. Arg., while
Phaeographis subtricosa (C. Knight) Miill. Arg., Phaeographis intumescens Miill. Arg. and Phaeographis
extenuata are taxonomic synonyms of Phaeographis subintricata Miill. Arg. Lectotypes are selected
for Phaeographis australiensis Miill. Arg. and Graphis aulacothecia C. Knight. The ascospores in
Phaeographis pseudomelana Miill. Arg. are 4-locular and not 6-locular as reported in the protologue.
Introduction
The lichen family Graphidaceae (the so-called Script Lichens) includes a number of
genera which are separated by the structure and colour of the ascospores, the presence
or absence of lirellae immersed in stromatic tissue and the structure of the paraphyses.
The distinction between these genera is best shown in a simplified key.
Key to the Australian Graphidaceae
1. Ascospores septate. 2
Ascospores muriform . 6
2. Ascospores hyaline. 3
Ascospores brown . 5
3. Lirellae scattered . 4
Lirellae grouped in stromatic tissue . Glyphis
4. Paraphyses simple . Graphis
Paraphyses branched . Diplogramma
5. Lirellae grouped in stromatic tissue . Sarcographa
Lirellae scattered . Phaeographis
6. Ascospores brown. 7
Ascospores hyaline. 8
462
Telopea 8 ( 4 ): 2000
7. Lirellae grouped in stromatic tissue . Sarcographina
Lirellae single . Phaeographina
8. Lirellae grouped in stromatic tissue . Medusulina
Lirellae single . 9
9. Ascospores 1-8 per ascus; paraphyses simple . Graphina
Ascospores 1 per ascus; paraphyses branched. Cyclographina
Apart from Glyphis (Acharius 1814), Snrcographa (Fee 1824) and Cyclographina (Awasthi
& Joshi 1979), the remaining genera were described by Muller: Graphina (1880),
Phaeograpliis (1882a: 336), Phaeographina (1882a: 398), Sarcographina (1887b),
Diplogramma (1891b) and Medusulina (1894).
The genera Phaeograpliis and Phaeographina were based on ascospore colour and
structure only and were not universally accepted. Vainio, in his account of the lichen
flora of Brazil (Vainio 1890), relegated them to subgenera in the genus Graphis and this
arrangement was followed in his account of the Philippine Graphidaceae (1920). Later,
however, botanists dealing with the Graphidaceae from the United States (Fink 1935),
Brazil (Redinger 1933, 1935), Mexico (Wirth & Hale 1963), New Zealand (Hayward
1977), Dominica (Wirth & Hale 1978), the United Kingdom (Purvis et al. 1992) and
Australia (Rogers & Hafellner 1992) have accepted Muller's concept of the genera
which has also been retained in the latest edition of the Dictionary of the Fungi
(Hawksworth et al. 1995). A detailed description of each of the two genera is given by
Rogers (1981).
Wirth and Hale (1978) discussed generic separation in the Graphidaceae and gave
examples of borderline species which could be placed in either of two genera, e.g.
Graphis or Graphina. They indicated the disadvantages of rejecting the spore-based
genera which would, in their view, leave only two choices, viz: the creation of many
more smaller genera with more closely related species, or the reduction of all the
spore-based genera back into the genus Graphis, which contains over 1000 species. In
the light of these difficulties they accepted, faute de mieux, the spore-based genera on
the grounds of practicality.
The family Graphidaceae is predominantly tropical to subtropical with relatively
fewer species found in temperate regions. For example only four Phaeograpliis species
(and no Phaeographina species) are reported from Great Britain (Purvis 1992) in contrast
to a total of 54 Phaeograpliis and Phaeographina taxa reported from Brazil (Redinger
1933) and 30 (including several unnamed species) from Australia.
The lirellae in Phaeograpliis and Phaeographina are often large and conspicuous but may
be immersed and inconspicuous. Ascospores range from c. 20 pm long and 4-locular
to c. 200 pm long and densely muriform. Lichen compounds are often absent but
species may contain norstictic or stictic acids (common in tire genera Graphis and
Graphina) and recently nomotatic, hypoprotocetraric, echinocarpic and hypostictic
acids were found in species of Phaeograpliis and Phaeographina from Australia and
elsewhere (Archer & Elix 1999).
As a number of new species and new reports remain to be published, a fuller
treatment, with keys, is not provided here.
Archer, Phaeographis and Phaeographina
463
Material and methods
The material examined consisted of type and other specimens from BRI, G, NSW and
WELT and additional recent collections from CANB, ESS, GZU, herb. Hafellner, herb.
Kalb and the author's own collections. Tire techniques used were described previously
(Archer 1999) but cross sections of lirellae were drawn in the dry state and were later
examined in water to determine the colour of the ascospores. Specimens were
photographed at a magnification of 6.5 on Fujichrome ASA 100 film and converted to
black and white, to give a final magnification of 17.
Phaeographis species
Phaeographis australiensis Mull. Arg. (Fig. la, 2a)
(Muller 1882b: 504).
Lectotype (here selected): New South Wales: Parramatta, W. Woolls s.n. (G).
Phaeographis subcompulsa Mull. Arg.
(Muller 1882b: 503).
Type: New South Wales: Nepean River [Sydney], W. Woolls s.n., 1880 (holo G; iso MEL
515586).
Phaeographis cinerascens Miill. Arg.
(Muller 1882b: 503).
Type: New South Wales: Nepean River [near Sydney], W. Woolls 81 (holo G; iso MEL
515594).
Phaeographis inscripta Mull. Arg.
(Muller 1882b: 504).
Type: New South Wales: Nepean River [near Sydney], W. Woolls s.n. (holo G).
Graphis aulacothecia C. Knight
(C. Knight 1882: 41).
Lectotype (here selected): New South Wales: [near Sydney], C. Knight 7, Vol. 204, p. 7,
1880 (WELT); syntype: C. Knight 46, Vol. 204, p. 7,1880 (WELT).
Thallus pale greyish white, thin, corticolous, surface slightly cracked, smooth and dull;
apothecia lirelliform, conspicuous, black, numerous, straight or sinuous, sometimes
branched, subimmersed to sessile, l-3(-6) mm long, (0.15-)0.2-0.3 mm wide, lips
closed or becoming slightly open, smooth, usually with a well-defined, but sometimes
evanescent, thalline margin; proper exciple laterally or almost completely carbonised;
hymenium (100-)120-150 pm tall; ascospores 8 per ascus, 2-seriate, fusiform, pale
brown, (30-)35-45(-55) pm long, (5—)7—10 pm wide, (7-)8-10(-ll)-locular.
Chemistry: norstictic acid.
Also examined (12 out of 78): Queensland: sine loc., Bailey 109, no date (BRI, AQ 720200); Woolston,
S. Brisbane, Wilson 1636, Aug 1889 (NSW).
New South Wales: Border Ranges National Park, Lumbsch 11007c, Oct 1994 (ESS); Old Macleay
River Estuary, Stuarts Point, Elix 21348, Jan 1987 (CANB); Gosford, Erina Creek, Elix 4701, May 1978
(CANB); 2 miles [3 km] N of Dural, Briggs L 586, Feb 1964 (NSW); Panania, Kelso Creek, Pinner
29, May 1998 (NSW); Mongarlowe River, 20 km SE of Braidwood, Streimann 36735, May 1986
(CANB).
464
Tetopea 8 ( 4 ): 2000
a
Fig.l. Cross-sections of lirellae and ascospores. a, Phaeographis australiensis; b, P. elaeina;
c, P. eludens; d, P. necopinata; e, P. nornotatica; f, P. pseudomelana; g, P. subintricata. lirellae:
scale bar = 200 pm; ascospores: scale bar = 20 pm.
Archer, Phaeographis and Phaeographina
465
Victoria: Cunningham, Wilson 1494, Mar 1888 (NSW); Cunningham [Gippsland], Mario, along Cape
Conran Road, Verdon 4194, Nov 1978 (CANB).
Norfolk Island: Rocky Point Reserve, Streimann 31806, Dec 1984 (B, CANB).
Tasmania: Moores Hill, near Beaconsfield, Kantvilas 206/80, May 1980 (HO); Ringarooma Tier,
Kantvilns 47/95, July 1995 (HO).
Phaeographis australiensis is characterised by the normally closed, black, lirelliform
apothecia, the variably carbonised proper exciple and the presence of norstictic acid.
When the lips are open the species somewhat resembles the chemically similar Graphis
semiaperta Mull. Arg. but that species always has a completely carbonised proper
exciple and smaller ascospores. The ascospores in P. australiensis are usually terminally
rounded but may sometimes be terminally acute (Hayward 1977: 574, Fig. 7; Fig. la).
The younger ascospores are often hyaline and older ascospores are conspicuously
brown and shrivelled; on occasion, few mature brown ascospores may be seen.
Phaeographis australiensis is a common species which occurs in eastern Australia from
Queensland to Tasmania and appears to be particularly abundant around Sydney. It
occurs on a variety of substrates including species of Avicennia, Banksia, Cassia,
Casuaritia, Erythrina, Leptospermum, Ligustrum, Hakea, Mains, Melia, Persoonia, Pultenaea
and Telopea. It also occurs on Norfolk Island and is reported from New Zealand
(Hayward 1977).
Two syntypes of P. australiensis were seen. One of these is labelled 'LBn 533', a
reference to Muller's publication in Flora, Lichenologische Beitrag no. 533, and is
therefore selected as Iectotype.
The type material of Graphis aulacothecia from WELT consists of 2 syntypes, Knight 7
and Knight 46. Of these, Knight 7 is the larger of the two specimens and is here selected
as Iectotype as the other smaller specimen has very few apothecia. Graphis aulacothecia
was previously reported as a synonym of Phaeographis australiensis (Archer 1999).
The morphological variation found in the types of the five names cited above falls
within the range of variation seen in the many recent collections. The four later names
are based on material collected within a small area near Sydney and it is therefore not
surprising that the 11 specimens belong to the same species.
Knight reported brown ascospores to be present in Graphis aulacothecia, and also in
Graphis elaeina, G. subintricata and G. s ublricosa (Knight 1882) but Muller transferred
only the last three species to his new genus Phaeographis.
Phaeographis australiensis somewhat resembles R mucromta (Stirt.) Zahlbr., according to
the description given by Stirton (Stirton 1876), but the two species appear to be
distinct. Phaeographis australiensis occurs predominantly in eastern New South Wales
whereas P. mucronata was collected in the Riverina District in the west of the State. In
addition, P. mucronata was reported to give no colour with alkali (K-ve) (Stirton, loc.
cit.) suggesting the absence of norstictic acid, which is present in P. australiensis.
The chemistry of P. mucronata has not been reported and the type was not available for
examination.
Phaeographis elaeina (C. Knight) Mull. Arg. (Fig. lb, 2b)
(Muller 1895: 321).
Graphis elaeina C. Knight
(Knight 1882: 41).
Type: New South Wales: [near Sydney], C. Knight 51 (holo G).
466
Telopea 8(4): 2000
Fig. 2 a, Phaeographis australiensis Mull. Arg., lectotype (G); b, Phaeographis elaeina (C. Knight) MAIL
Arg., holotype (G); c, Phaeographis eludens (Stirt.) Shirley, Elix 35177 (CANB); d, Phaeograpliis
ne&pimta A -W-Archer & Elix, holotype (CANB); e, Phaeograpliis nomolalica A.W. Archer & Elix,
holotype (CANB); f, Phaeographis pseudomelana Mull. Arg., holotype (G). All xl7.
Archer, Phaeographis and Phaeographina
467
Thallus off-white to pale olive green, thin, corticolous, surface smooth and shiny;
apothecia lirelliform, black, conspicuous, immersed or sometimes subsessiJe, straight,
curved or sinuous, sometimes branched, 1-4 mm long, 0.2-0.5 mm wide, often in
substellate clusters to 4 mm wide, with a thin, white thalline margin; proper exciple
thin, completely carbonised; hymenium (75-)100-125 pm tall; disc matt black,
epruinose; ascospores 8 per ascus, irregularly 2-seriate, very pale brown,
(15-)17-21(-23) pm long, 6-7 pm wide, 4-locular.
Chemistry: no compounds found.
Also examined: Queensland: New England Highway, 20 km SW of Yarraman, Haffellner 19362, Sep
1986 (GZU); Conors Range, c. 20 km SW of Sarina, Lumbsch 10983d, p.p., Oct 1994 (ESS); Bowenia
State Forest, Stony Creek, 25 km NNW of Yeppoon, Elix 34571, Aug 1993 (CANB); Forrest Beach,
16 km SE of Ingham, Streinmnn 28849, June 1984 (CANB); Mcllwraith Range, 29 km NE of Coen,
Streitnann 56831, Oct 1995 (CANB).
New South Wales: Ku-ring-gai Chase National Park, the Basin camping area, on felled tree. Archer
G261, Sep 1998, (NSW).
Phaeographis elaeina is characterised by the immersed, relatively wide, black lirellae, the
4-locular ascospores and the thin, completely carbonised proper exciple. This latter
feature distinguishes the species from the somewhat similar P. subintricata, which has
a thin carbonised proper exciple open at the base (laterally carbonised), and
P. pseudomelana, which is apically carbonised. Phaeographis elaeina has been found in
Queensland and New South Wales. Reported substrates include Casuarina and
Excoecaria.
Phaeographis eludens (Stirt.) Shirley (Fig. lc, 2c)
(Shirley 1889: 197).
Graphis eludens Stirt.
(Stirton 1881: 72).
Type: Queensland: sine loc., as 'saxicola', F. Bailey 287 (holo BRI).
Thallus fawn to pale fawn, 1-2 mm thick, saxicolous, surface smooth and dull
(superficially resembling fine velvet); apothecia lirelliform, black, numerous,
inconspicuous, open, immersed, straight, curved or sinuous, often branched and in
sub-stellate clusters; individual lirellae 0.5-2 mm long, 0.1-0.2 mm wide, lacking a
thalline margin; proper exciple absent; hymenium (100—)125—175(—200) pm tall;
ascospores 8/ascus, 2-3-seriate, pale brown, 12-15 pm long, 5-7 pm wide, 4-locular.
Chemistry: no compounds found.
Also examined: Queensland: sine loc., Shirley s.n., Jan 1888 (NSW); Burleigh Heads National Park,
on basalt boulders, Elix 1093, Aug 1975 (CANB); Isla Gorge National Park, 27 km NNE of Taroom,
on sandstone rocks, Elix 35177, Aug 1993 (CANB); Tabletop, Harvey Range, 39 km WSW of
Townsville, Streimann 37185, 37192, Aug 1986 (CANB).
Phaeographis eludens is characterised by the substrate preference, the thick thallus, the
small 4-locular ascospores and the absence of lichen compounds. Stirton (1881)
compared his new species with Graphis hypoglauca Krempelh. [Phaeographis hypoglauca
(Krempelh.) Zahlbr.] but the ascospores in that species are larger, 18-28 x 8-11 pm
(Singh & Awasthi 1979).
The species is so far known only from Queensland where it occurs on basalt and
sandstone. The species was also reported from Moggill and Mount Perry [Brisbane]
(Shirley 1889).
468
Telopea 8(4): 2000
An unnamed saxicolous species of Phaeogrciphis from Queensland with 4-locular
ascospores (H. Streimann 56404, CANB) is differentiated from P. eludens by the
tuberculate thallus and the conspicuous thalline margins of the lirellae.
Phaeographis necopinata A.W. Archer & Elix (Fig. Id, 2d)
(Archer & Elix 1999: 92).
Type: Queensland: Great Dividing Range, Mount Baldy, 4 km S of Atherton, on
Alphitonia, J.A. Elix 16233, June 1984 (holo CANB).
Thallus off-white, thin, corticolous, surface smooth and shiny; apothecia lirelliform;
lirellae numerous, conspicuous, sessile, open, straight, curved or sinuous, sometimes
in substellate clusters, with a thin thalline margin, 1-3 mm long, 0.3-0.5 mm wide, and
clusters to 4 mm wide; proper exciple completely carbonised; hymenium 125-150 pm
tall; epithecium matt black, epruinose or lightly pruinose; ascospores pale brown to
brown, 8 per ascus, (25-)30-40 pm long, (8-)9-12 pm wide, (7-)8-locular.
Chemistry: hypostictic acid (major), conhypoprotocetraric acid (trace) and
hyposalazinic acid (trace).
Also examined: Queensland: Lamb Range, 21 km NE of Atherton, Streimann 29822, June 1984
(CANB); Bellenden Ker, Mayrhofer 11921, Aug 1993 (GZU); type locality, Elix 16275, June 1984
(CANB).
Phaeographis necopinata is characterised by the conspicuous lirellae, asci with eight
ascospores and the presence of hypostictic acid. The lirellae may be terminally
rounded or acute and the ascospores are predominantly 8-locular. Superficially the
new species resembles Phaeographis dendritica (Ach.) Mull. Arg., but is distinguished
from that species by the presence of hypostictic acid. Hypostictic acid is an uncommon
compound in the Graphidaceae but is also found in two species from Brazil, viz. the
holotype of Graphina albostriata (Vain.) Zahlbr. (TUR-V 27177) and a syntype of
Graphina pseudosophisticata (Vain.) Mull. Arg. (TUR-V 27244A) (Archer & Elix 1999).
These two taxa are conspecific.
Phaeographis nornotatica A.W. Archer & Elix (Fig. le, 2e)
(Archer & Elix 1999: 93).
Type: Queensland: Mcllwraith Range, Llankelly Creek, 9 km NE of Coen, H. Streimann
56937, Oct 1995 (holo CANB).
Thallus dull fawn, thin, corticolous, surface smooth and shiny (chondroid); apothecia
lirelliform, conspicuous, scattered, sessile, black, straight, curved or sinuous,
sometimes branched, with a conspicuous thalline margin, lips open, disc black, fine
white-pruinose, l-4(-6) mm long, 0.5-0.7(-0.9) mm wide, 0.4-0.5 mm tall; proper
exciple completely carbonised, laterally thin, the base thick, c. 250 pm tall, tapering
towards the base; hymenium c. 100 pm tall; ascospores 8 per ascus, pale brown,
rounded cylindrical, 21-25 pm long, 6-7 pm wide, 6-locular.
Chemistry: 4-O-demethylnotatic acid [nornotatic acid] (major) and hypoprotocetraric
acid (minor).
1984 (CAr4B) Cd: Queensland: Bi ® Tabieland ' 26 km S of Cooktown, on Alphitonia, Elix 17243, July
Phaeographis nornotatica is characterised by the large conspicuous lirellae, the tapering
base of the proper exciple and the presence of nornotatic acid. The species somewhat
resembles P. exaltata (Mont. & v.d. Bosch) Mull. Arg. but the lirellae are larger and
ta er, and the proper exciple and the chemistry differ. Phaeographis exaltata lacks lichen
compounds. The new species is so far known only from the two specimens cited above.
Archer, Phaeographis and Phaeographina
469
Phaeographis pseudomelana Mull. Arg. (Fig. If, 2f)
(Muller 1895: 321).
Type: Queensland: sine loc., /. Shirley 1838, 1893 (holo G).
Thallus pale fawn, thin, corticolous, surface smooth and shiny; apothecia lirelliform,
conspicuous, scattered, sessile, open, straight, curved or sinuous, often branched,
terminally acute, with a thin thalline margin, 2-4(-6) mm long, 0.15-0.25 mm wide;
proper exciple apically carbonised; hymenium 100-125 pm tall; disc black, white-
pruinose; ascospores 8 per ascus imbricate 1-seriate, pale brown, 17-20 pm long, 6-8
pm wide, 4-locular.
Chemistry: no compounds found.
Also examined: Queensland: Rainbow Beach, c. 50 km W of Gympie, Lumbsch 10995j, Oct 1994
(ESS); Kirrima State Forest, 24 km WNW of Cardwell, Elix 15685, June 1984 (CANB).
New South Wales: Monga State Forest, J. Everett 947, May 1986 (NSW).
Phaeographis pseudomelana is characterised by the open lirellae, the apically carbonised
proper exciple, the 4-locular ascospores and the absence of lichen compounds. It
closely resembles P. subintricata (C. Knight) Mull. Arg. and is only distinguished from
that species by the apically carbonised proper exciple, in contrast to the laterally
carbonised proper exciple in the latter species. The type and other specimens
examined possess 4-locular ascospores in agreement with Muller's drawing in sched.
although the protologue refers to 6-locular ascospores. Shirley (1896) reported the
species from Sankeys Scrub [Brisbane].
Phaeographis subintricata (C. Knight) Mull. Arg. (Fig. lg, 4a)
(Muller 1895: 320).
Graphis subintricata C. Knight
(Knight 1882: 40).
Type: New South Wales: [near Sydney], C. Knight 41 (syntype WELT); additional
syntype: G.
Phaeographis subtricosa (C. Knight) Mull. Arg.
(Muller 1895: 320).
Graphis subtricosa C. Knight
(Knight 1882: 40).
Type: New South Wales: [near Sydney], C. Knight (Vol. 69A, p.15), no. 6 (holo WELT;
iso G).
Phaeographis intumescens Mull. Arg.
(Muller 1893: 56).
Type: Victoria: Lakes Entrance, F.R.M. Wilson 877 (holo G).
Phaeographis extenuata Mull. Arg.
(Muller 1893: 57).
Type: Victoria: Lakes Entrance, F.R.M. Wilson 877 (holo G).
Thallus off-white to pale olive green, thin, corticolous, surface smooth and slightly
shiny; apothecia lirelliform, black, conspicuous, open, immersed, becoming subsessile,
470
Telopea 8(4): 2000
straight, curved or sinuous, often branched, 1-4 mm long, 0.2-0.5 mm wide, often with
a conspicuous, white thalline margin, sometimes forming substellate clusters; proper
exciple thin, laterally carbonised, open at the base; hymenium 100-125 pm tall; disc
matt black, epruinose; ascospores 8 per ascus, irregular 2-seriate, very pale brown,
(14-)17-21(-24) pm long, 6-8 pm wide, 4-locular.
Chemistry: no compounds found.
Also examined: Queensland: Upper Coomera, Wilson s.n., as 'Graphis innata' (NSW).
New South Wales: Sydney, Mosman Bay, Wilson s.n., Sep 1897 (NSW); Urunga, Hungry Head,
c. 25 km SSW of Coffs Harbour, Archer G 212, Apr 1998 (NSW).
Victoria: Cunningham, Wilson s.n., no date (NSW L5082); Metung, Wilson 877a, Mar 1889 (NSW);
Lake Tyers, Wilson, no date (NSW 180877); Metung, Wilson s.n., Mar 1889 (NSW); Gippsland, 7 km
S of Bonang, Verdon 4169, Nov 1978 (CANB, H).
New Zealand: sine loc., C. Knight s.n. [as Graphis In/poleuca, nom. nud.] (NSW 155588).
Phaeographis subintricata is characterised by the black, open lirellae, 4-locular
ascospores, the incompletely carbonised proper exciple and the absence of lichen
compounds. It resembles P. dendritica but is distinguished from that species by the
smaller, 4-locular ascospores and the absence of norstictic acid. Phaeographis
subintricata closely resembles P. elaeina and is only distinguished from that species by
the completely carbonised proper exciple in the latter species, and from
P. pseudomelana by the apically carbonised proper exciple in that species.
Graphis subintricata and Graphis subtricosa were published simultaneously (Knight
1882) and as G. subintricata is the first Graphis species described (no. 5; G. subtricosa no.
6) it is taken to be the earliest name. Phaeographis intuntescens and P. extenuata were also
published together (Muller 1893, loc. cit.) and are based on the same specimen, Wilson
877, but the two new species were differentiated on the basis of the thallus colour.
Muller (loc. cit., 1895) noted that P. subintricata was close to P. extenuata. The four
species listed lack any features to distinguish between them.
Reported substrates include Acacia, Eucn/phia, Lomatia and Notelaea.
Additional syntypes of P. subintricata are in G, M and WELT (Filson 1986) and as all of
these have not been seen, no lectotype is selected here.
Species of uncertain status
Phaeographis mucronata (Stirt.) Zahlbr.
(Zahlbruckner 1923: 382)
Graphis mucronata Stirt.
(Stirton 1876: 95).
Lectotype (fide Rogers 1982): New South Wales: Riverina District, H. Paton s.n., 1875
(BM, not seen).
The type was not available for examination; see discussion above under Phaeographis
australiensis.
Archer, Phaeographis and Phaeographina
471
Phaeographina species
Phaeographina banksiae Mull. Arg. (Fig.3a, 4b)
(Muller 1893: 59).
Type: Victoria: Maffra, on Hymenanthera banksii, F.R.M. Wilson 879, 1892 (holo G; iso
NSW).
Thallus pale reddish-brown, thin, corticolous, surface smooth and slightly shiny;
apothecia lirelliform, black, numerous, conspicuous, sessile to slightly immersed, lips
closed, becoming slightly open, straight or curved, rarely branched, 1-3 mm long,
0.2-0.3 mm wide, with a thin, evanescent thalline margin; proper exciple laterally
carbonised; hymenium 125-150 pm tall; ascospores 8 per ascus, pale brown, ellipsoid,
(20-)24-31(-39) x (10-)12-15(-17) pm, 4-6 x 2-3-locular.
Chemistry: no compounds found.
Also examined: Queensland: Upper Coomera, Wilson 1547, no date (NSW).
New South Wales: Jenolan Caves, Wilson s.n., Sep 1897 (NSW).
Fig. 3. Cross-sections of lirellae and ascospores. a, Phaeographina banksiae; b, P. echinocarpica;
c, P. muelleri, lirellae: scale bar = 200 pm; ascospores: P. banksiae: scale bar = 20 pm, others:
scale bar = 100 pm.
472
Telopea 8(4): 2000
Victoria: Maffra, Wilson s.n. (NSW L5084); Sale, on Grevillea robusta, Wilson 879, Sep 1886 (NSW);
bank of Yarra River, Kew, Wilson s.n., Dec 1884 (NSW).
Phaeographina banksiae is characterised by the black lirellae, the laterally carbonised
proper exciple, the small muriform ascospores and the absence of lichen compounds.
It is distinguished from other Australian Plweographina species by the small
ascospores. The species is so far known from Queensland, New South Wales and
Victoria.
Phaeographina echinocarpica A.W. Archer & Elix (Fig. 3b, 4c)
(Archer & Elix 1999: 91).
Type: Queensland: Mount Baldy, 4 km SW of Atherton, 17°17'S, 145°27’E, alt. 1080 m,
in old Araucaria plantation and regrowth rainforest, on Alphitonia, J.A. Elix 16275, June
1984 (holo CANB).
Fig. 4. a, Phacographis subiiitricata (C. Knight) Mull. Arg., syntype (G); b, Phaeographina banksiae Mull.
rg holotype (Gp, c, Phaeographina echinocarpica A.W. Archer & Elix, holotype (CANB);
d, Phaeographina muelleri A.W. Archer, holotype (G). All xl7. ^
Archer, Phaeographis and Phaeographina
473
Thallus pale olive green, thin, corticolous, surface smooth and shiny. Lirellae
conspicuous, numerous, scattered, sessile, greyish black, with conspicuous thalline
margin, curved or sinuous, sometimes branched, 1-7 mm long, 0.7-0.9(-1.0) mm wide;
proper exciple laterally carbonised, red-brown at the base; hymenium 200-250 pm tall;
ascospores 1 per ascus, pale brown, elongate ellipsoid, 162-200 pm long, 30-40 pm
wide, muriform (c. 20 x 5-6 locular).
Chemistry: echinocarpic acid (major) and 4-O-methylechinocarpic acid
[conechinocarpic acid] (minor).
Also examined: Queensland: Cardwell Range, 46 km SE of Ravenshoe, Elix 16117 p.p, June 1984
(CANB).
Phaeographina echinocarpica is characterised by the large, pale brown ascospores, the
conspicuous lirellae and the presence of echinocarpic acid. The carbonised proper
exciple is covered with a thin thalline layer thus giving the lirellae a greyish black
appearance.
Echinocarpic acid is an uncommon benzyl ester distinguished from other esters of this
type, such as alectorialic acid and barbatolic acid, by ester formation involving an
ortho- rather than a meto-hydroxymethyl group in the B-ring (Elix et al. 1995).
Previously echinocarpic acid had been reported only from the Parmeliaceae.
The new species is distinguished from other species of Phaeographina with large
ascospores, such as P. montagnei (v.d. Bosch) Miill. Arg. and P. muelleri (vide infra), by
the presence of echinocarpic acid.
The species is so far known only from the two specimens cited.
Phaeographina muelleri A.W. Archer, sp. nov. (Fig. 3c, 4d)
Synonym: Phaeographina caesiopruinosa (Fee) Miill. Arg. var. monospora Miill. Arg., Bull.
Herb. Boissier 3: 322 (1895).
Type: Queensland: Toowoomba, C. Hartmann s.n. (holo G).
Thallus pale fawn, thin, corticolous, surface smooth and shiny; apothecia lirelliform,
conspicuous, scattered, conspicuously sessile, open, with a conspicuous thalline
margin, 1-2 mm long, 0.3-0.7 mm wide; proper exciple apically carbonised;
hymenium 175-200 pm tall; disc black, white-pruinose; ascospores 1 per ascus,
elongate-ellipsoid, pale brown, 145-180 pm long, 30-40 pm wide, densely muriform.
Chemistry: no compounds found.
Phaeographina muelleri is characterised by the conspicuous, open, apically carbonised
lirellae, the large muriform ascospores and the absence of lichen compounds. It is
distinct from other species of Phaeographina with large ascospores and lacking lichen
compounds which occur in Australia such as P. montagnei (v.d. Bosch) Miill. Arg.,
which has a bright red epithecium and a completely carbonised proper exciple, and an
unnamed species from Queensland which has closed lirellae and lacks a carbonised
proper exciple.
Phaeographina muelleri is so far known only from the type specimen.
Phaeographina caesiopruinosa (Fee) Miill. Arg. (Miiller 1887a) is reported to be a
common New World taxon with brown muriform ascospores, 4-8 per ascus, 45-120 x
15-20 pm and lacking lichen compounds (Miiller 1894; Wirth & Hale 1978). Muller
(1887a) reported the taxon to occur in Ceylon and Java but it was not listed by
Leighton from Ceylon (Leighton 1869) nor from Indonesia by Redinger (1936) and in
addition it was not reported from adjacent areas such as Japan (Nakarvishi 1966,1977)
and the Philippines (Vainio 1920). The taxon was reported from Queensland by Shirley
474
Telopea 8(4): 2000
(1891) and Muller (1891a, 1891b) but no specimens of the taxon were found in a recent
examination by the author of Shirley and Bailey specimens from BRI and it has not
been found among the many recent collections from Queensland in CANB. It is
possible that the specimens examined were Phaeographina quassiicola (Fee) Mull. Arg.,
which has ascospores similar to those of P. caesiopruinosa. Phaeographina caesiopruinosa
thus remains a New World species, confirmed by recent collections from Mexico
(Wirth & Hale 1963) and Dominica (Wirth & Hale 1978). As var. caesiopruinosa appears
to be absent from the Australian region and the Australian taxon is distinct from var.
caesiopruinosa, in that it has an apically carbonised exciple and asci with one large
ascospore, specific status is appropriate for the local Australian variety which is here
named Phaeographina muelleri. The epithet commemorates the Swiss lichenologist Jean
Muller of Argau (Mull. Arg.), 1826-1896, who described a large number of lichen taxa
from Australia, including the present taxon.
Acknowledgments
The author is grateful to the Herbaria cited above for the loan of type and other
specimens and to the National Herbarium of New South Wales (NSW) for permission
to use the facilities of the Herbarium, for arranging the loan of the specimens cited
above and to Ms. C. Wardrop (NSW) for the drawings.
References
Acharius, E. (1814) Glyphis. Syn. Lick: 106 (Svandborg et Soc.: Lund).
Archer, A.W. (1999) The lichen genera Graphis and Graphina (Graphidaceae) in Australia 1. Species
based on Australian type specimens. Telopea 8: 273-295.
Archer, A.W. & Elix, J.A. (1999) Three new species in the Australian Graphidaceae with novel
chemistries: Phaeographina echinocarpica, Phaeographis necopinata and Phaeographis nomotatica.
Mycotaxon 72: 91- 96.
Awasthi, D.D. & joshi, M. (1979) The lichen genera Helminlhocarpon, Cyclographa and Cyclographina
(gen. nov.). Norzveg. J. Bot. 26:165-177.
Elix, J. A., Lajide, L. & Wardlaw, J.H. (1995) The structure of echinocarpic acid. A benzyl ester from
Pamtelia norcrambidocarpa. Austral. /. Chew. 48:1213-1216.
Filson, R.B. (1986) Index to type specimens of Australian lichens 1800-1984. Australian Flora and
Fauna Series Number 4 (Australian Government Publishing Service: Canberra).
Fink, B. (1935) The Lichen Flora of the United States: 106-119 (The University of Michigan Press:
Ann Arbor).
Hawksworth, D.J., Kirk, P.M., Sutton, B.C. & Pegler, D.N. (1995) Dictionary of the Fungi. (University
Press: Cambridge).
I layward, G.C. (1977) Taxonomy of the lichen families Graphidaceae and Opegraphaceae in New
Zealand. Nero Zealand J. Bot. 15: 565-584.
Knight, C. (1882) Contributions to the Lichenographia of New South Wales. Trans. Linn. Soc. London,
Bot . 2: 37-51.
Leighton, W.A. (1869) The Lichens of Ceylon. Trans. Linn. Soc. Lond. (Bot) 27:161-185.
Muller, J. (1880) Lichenologische Beitrage X. Flora 63:17-22.
Muller, J. (1882a) Lichenologische Beitrage XV. Flora 65: 326-337, 397-402.
Muller, J. (1882b) Lichenologische Beitrage XVI. Flora 65: 483-490,499-505.
Muller, J. (1887a) Graphideae Feeanae. Mem. Soc. Phys. Hist. Nat. de Geneve 29:1-80.
Muller, J. (1887b) Lichenologische Beitrage XXVI. Flora 70: 423-429.
Muller, J. (1891a) Lichenes Bellendenici. Hedivigia 30: 47-56.
Muller, J. (1891b) Lichenes Brisbancnses. Niton Giorn. Bot. Hal. 23: 385-404
Muller, J. (1893) Lichenes Wilsoniani. Bull. Herb. Boissier 1: 33-65.
™ u " eT ' I' ir ichenes Eckfcld tiani in Louisiana et Florida. Bull. Herb. Boissier 2: 41-50; 89-93.
Muller, J. (1895) Sertum Australiense. Bull. Herb. Boissier 3: 313-327.
Archer, Phaeographis and Phaeographina
415
Nakanishi, M (1966) Taxonomkal Studies on the family Graphidaceae of Japan. ]. Sci. Hiroshima
Unit’., Ser. B, Div. 2, Bot. 11:1-126.
Nakanishi, M. (1977) Notes on Japanese species of Phaeographina. Hikobia 8: 91-100.
Purvis, O.W., Coppins, B.J., Hawksworth, D.J., James, P.W., & Moore, D.M. (1992) The Lichen Flora
of Great Britain and Ireland. (Natural History Museum Publications: London).
Redinger, K.M. (1933) Die Graphideen der ersten Regnell'schen Expedition nach Brasilien,
1892-1894. II: Graphina und Phaeographina. Ark. Bot. 26A(1): 1-105.
Redinger, K.M. (1935) Die Graphideen der ersten Regnell'schen Expedition nach Brasilien,
1892-1894. Ill: Graphis und Phaeographis. Ark. Bot. 27A(3): 1-103.
Redinger, K.M. (1936) Die Graphideen der Sunda Inseln. Rev. Bryol. Lichenol. 9: 37-122.
Rogers, R.W. (1981) The Genera of Australian lichens (Lichenised Fungi). (University of Queensland
Press: St. Lucia).
Rogers, R.W. (1982) Typification of the species of lichens described from Australian specimens by
James Stirton. Austrobaileya 1: 502-510.
Rogers, R.W & Hafellner, J. (1992) A systematic arrangement of the Australian lichens. Flora of
Australia 54: 46-65.
Shirley, |. (1889) The lichen Flora of Queensland. Proc. Roy. Soc. Queensland 6:165-218.
Shirley, J. (1891) Lichenes. In Bailey, F.M., Contributions to the Queensland Flora. Queensland Dept.
Agric. Bulletin 9, Botany Bulletin s.n. [3:] 29-30.
Shirley, J. (1896) Lichenes. In Bailey, F.M., Contributions to the Queensland Flora. Queensland Dept,
of Agric. Bulletin s.n., Botany Bulletin XIII: 27-28.
Singh, K.P. & Awasthi, D.D. (1979) Lichen genus Phaeographis from India and Ceylon. Bull. Bot. Sum
India 21: 97-120.
Stirton, J. (1876) Lichens British and Foreign. Trans. Glasgow Field Naturalists 4: 85-95.
Stirton, J. (1881) Additions to the lichen Flora of Queensland. Trans & Proc. Roy. Soc. Victoria
17: 66-78.
Vainio, E.A. (1890) /tude sur la classification naturelle et la morphologie des Lichens du Bresil. Acta
Soc. Fauna FI. Fenn. 7:1-247.
Vainio, E.A. (1920) Lichenes insularum Philippinarum 111. Ann. Acad. Sci. Fenn., Ser. A. 15,6:1-368.
Wirth, M. & Hale M.E. (1963) The lichen family Graphidaceae in Mexico. Contr. U. S. Natl. Herb.
36: 63-119.
Wirth, M. & Hale M.E. (1978) Morden-Smithsonian Expedition to Dominica: The Lichens
(Graphidaceae). Smithsonian Contr. Botany.: 1-64.
Zahlbruckner, A. (1923) Catalogus lichenum universalis, vol. 2. (Borntraeger: Leipzig).
Manuscript received 2 September 1999
Manuscript accepted 15 May 2000
mm
mm.
,
■
,
477
Hopkinsiaceae and Lyginiaceae, two new
families of Poales in Western Australia, with
revisions of Hopkinsia and Lyginia
Barbara G. Briggs and L.A.S. Johnsont
Abstract
Briggs, Barbara G. & Johnson, L.A.S. (Royal Botanic Cardens, Mrs Macquaries Road, Sydney, NSW 2000,
Australia) 2000. Hopkinsiaceae and Lyginiaceae, tivo new families of Poales in Western Australia, xoith
revisions o/Hopkinsia and Lyginia. Telopea 8 (4): 477-502. Hopkinsia and Lyginia are excluded from
Restionaceae and the new families Hopkinsiaceae and Lyginiaceae established. DNA sequence
data from the chloroplast gene rbcL, the trnL intron and tmL-trnF intergenic spacer indicate that
these genera are more closely allied to Anarthriaceae than to Restionaceae, although differing
markedly from the former in vegetative morphology' and anatomy. The contrasting results of
cladistic analyses of DNA sequence data and morphological data are discussed, as are tire reasons
for excluding these genera from Restionaceae and Anarthriaceae. Extensive descriptions of the new
families are given and a key' provided to these and related families. The features that they share
with Restionaceae are considered to be largely plesiomorphic within Poales or associated with their
occurrence, like many Restionaceae, in conditions of seasonal drought. Occurring in a region of
great diversity of Restionaceae, their distinctiveness was overlooked until the new DNA evidence
became available. Distinctive features for Hopkinsia include the ovary' structure, style with stigmatic
branches that are themselves branched, fleshy pedicels, and fruit with both woody and fleshy
layers. Distinctive for Lyginia are the oblique epidermal cells and stomates, distinctive
chlorenchyma structure, fused staminal filaments, seed ornamentation and presence of unusual
chemical constituents (allose, allosides and fructan-ty'pe oligosaccharides). Both genera also differ
from Restionaceae in having 2-lobed, 4-sporangiate anthers and pollen of graminoid ty'pe. Two new
species, H. adscendens from south coastal Western Australia and L. excelsa from tire Cataby region
north of Perth are described, and the differences in reproductive biology between the species of
Lyginia are noted. Both the species of Hopkinsia and L. adscendens have vulnerable or endangered
conservation status. The chromosome number 2 n = 18 is reported in H. adscendens.
Introduction
Although Hopkinsia and Lyginia have until now been included within Restionaceae,
they have long been recognised as differing substantially from other members of that
family. Gilg-Benedict (1930) placed them, with Anarthria and Ecdeiocolea, in
Diplantherae, characterised by bithecate anthers, whereas all other Restionaceae have
unithecate anthers and constituted the Haplantherae. Cutler and Shaw (1965)
removed Anarthria and Ecdeiocolea from Restionaceae establishing two new families. In
the absence of molecular data, we have long been aware of major differences between
Hopkinsia and Lyginia and all other genera then included in Restionaceae (Johnson
& Briggs 1981), although we regarded them as basal groups within that family. More
recently, since DNA data became available, we have foreshadowed recognition of the
new families here described (Linder et al. 1998; Briggs & Johnson 1999; Briggs et al.
2000; Briggs 2000).
t Deceased 1 August 1997.
478
Telopea 8(4): 2000
Evidence supporting the recognition of new families
Members of the same plant family can be expected to form a monophyletic group
separate from related families; they should also share synapomorphic features by
which the family can be characterised and contrasted with other families. Criteria of
relationship and morphology must both be met.
In the present case, in brief, molecular evidence requires the exclusion of Hopkinsia and
Lyginia from Restionaceae. Morphological evidence gives no basis for these genera
being placed in Anarthriaceae and similarly fails to give a basis for uniting them in a
single new family. The analyses of morphological and DNA data in this case produce
contradictory results, but these appear to indicate a degree of homoplasious and
convergent morphological evolution, rather than implying substantially different
phylogenies.
The justification for recognising the new families Hopkinsiaceae and Lyginiaceae
depends on establishing that:
(1) Hopkinsia and Lyginia are not correctly placed within the Restionaceae;
(2) synapomorphies shared with Anarthriaceae are not sufficient for them to be
included in that family; and
(3) they are not appropriately placed together in a single new family.
(1) Exclusion from Restionaceae
ie major evidence for exclusion of Hopkinsia and Lyginia from Restionaceae comes
rom analysis of DNA sequence data. Cladistic parsimony analysis of DNA sequences
° *■ e chloroplast gene rbcL and of the trn L intron, together with the tniL-lniF
intergenic spacer, does not associate Hopkinsia and Lyginia with other genera referred
to Restionaceae but shows a robust clade of these two genera with Anarthria
(Mg. 1 modified from Briggs et al. 2000). This clade has 100% jackknife support in
separate cladistic analyses of each of the two regions of the chloroplast DNA, as well
as ot the combined data. In addition, the clade is marked by two indels in the trnL
m on, an insertion of five bases and a deletion of three; these indels show no
tiomoplasy among other members of the Poales studied. This molecular evidence
comes w io y rom chloroplast DNA but is supported by the highly conserved rbcL
gene, as v\ e as a less conserved region of DNA that is now being widely used in
phylogenetic studies (e.g. Bayer & Starr 1998) and has given moderately well resolved
rees in our studies of Poalean families. The consensus trees from the combined DNA
r . f , aCeS , e ' ,r ' a c l fl de as sister to the Restionaceae clade (Restionaceae with
l . ? e ^! L 3C -?f e ' ®ta from rbcL alone similarly distinguish these and the Poaceae
tee eaciwi l ,, jackknife support) but places them (and several other families)
at an unresolved polychotomy.
t / le reSU ^j f' orn studies (Fig. 1) as giving a clear indication that
in a rMH Hopkt ™\ md h ^ mw form * clade. These genera, however, fail to associate
!IiIrf an !i‘ 1S , 0nnl0rph0lft8ic ' 11 data ( pi 8- 2 ' modified from Linder et al. 2000);
Restinmr ”‘rru ^‘ nm , a ' ld Hopkinsia emerge successively at the base of
Centrnlpn H 6 cladograms also show contrasting positions for the
noTdwiK?!>H ea ° f that iamii V are being further investigated and are
Dositions nf r m 10 P resent P a P er - Similarly, the two trees show differences in the
are beimr § en fra within Australian Restionaceae and these discrepancies
arise from h f U ' P ' ^. uc ' 1 a contr asting result for Hopkinsia and Lyginia could
perhans durinp 0 ^ .'T'r' 5 3n convergent change in morphological characters,
morDholn(rinl^ eVO U '° n L , n ? C f r s ‘ mi ' ar environmental conditions. Constraining the
P g c tree so that Hopkinsia and Lyginia associate with Anarthria, rather than
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
479
53
82
99
74 .
88
89 ,
82
53
100
80
80
75
97
81
96
100
HI
98
64
+t
100
I
64
70
95
94
64
93
88
H
100
H
100
H
67
100
99
100
- Desmocladus castaneus _
- Harperia lateriflora
- Kulinia eludens
- Coleocarya gracilis
- Lepidobolus chaetocephalus
- Dielsia stenostachya
- Baloskion gracile
- Baloskion tetraphyllum
- Melanostachya ustulata
_ Tyrbastes glaucescens
- Guringalia dimorpha
- Acion hookeri
- Saropsis fastigata
- Chordifex stenandrus
- Chordifex amblycoleus
- Chordifex jacksonii
- Tremulina tremula
- Loxocarya gigas
- Meeboldina coangustata
- Meeboldina cana
j
- Leptocarpus tenax
- Dapsilanthus ramosus
- Chaetanthus aristatus
- Alexgeorgea ganopoda
- Winifredia sola
_ Taraxis grossa
L_ Empodisma minus
- Eurychorda complanata
- Calorophus elongatus
- Lepyrodia glauca
- Sporadanthus tasmanlcus -
- Elegia cuspidata African Restlonaceae
- Centrolepis strigosa Centrolepidaceae
- Hopklnsia adscendens -
- Lyginia barbata
Anarthria
Clade
- Anarthria polyphylla
- Zea mays
- Triticum aestivum
- Oryza sativa
- Ecdeiocolea monostachya
- Georgeantha hexandra _
- Carex appressa/hostiana Cyperaceae
Poaceae
Clade
Fig. 1. Jackknife consensus tree from parsimony analysis of combined chloroplast DNA sequence
data (rbcL, IrnL intron and IrnL-trnf intergenic spacer) for representative species. Support indices
are shown and indels supporting clades are indicated by cross-bars; homoplasious indels are
shown by narrow bars and are identified by letters. The outgroup consists of a combination of the
rbcL sequence of Carex hostiana with the trriL—trnV region of C. nppressa. (Modified from
Briggs et al. 2000).
480
Telopea 8(4): 2000
with Restionaceae, increases tree length by four steps to 305 steps (Linder et al. 2000).
If, on the other hand, the molecular tree is constrained to match the morphological tree
by placing these genera as basal but ungrouped, tree length increases from 2742 steps
to 2807, an increase of 65 steps. In addition, both the two indels common to the three
genera would become homoplasious, requiring an additional four innovations.
Our morphological studies suggest that most of the characters that are common to
Restionaceae, Hopkfrish and Lyginia (Table 1), apart from dioecy and some aspects of
culm anatomy, are plesiomorphic within Poales or are readily interpreted as
homoplasies associated with occurrence in regions of low soil fertility and seasonal
drought. Indeed, within the Poales, one of the most conspicuous similarities shared by
Hopkinsia, Lyginia and Restionaceae is the xeromorphic habit with reduced leaf
laminae, so that the culms are the principal photosynthetic organs. Some further
relevant characters may arise from embryology (c.f. Linder & Rudall 1993) and from
inversions within the chloroplast DNA (Doyle et al. 1992), but information on these
features is lacking for Hopkinsia and Lyginia.
;=A!
Joinvillca
Ecdeoicolea
Gcorgcantlia
Anarthria
Hopkinsia <—
Lcpvrodia
Baloskion
Ac ion
Saropsis
Gunngalia
Choraifex
Eurvchorda
Diclsia
Trcniulina
Loxocarya
Cvtogomdium
Melanostachya
Platvchorda
Tv rbastcs
Gaimardia
Centrolepis
Brizula
Desmocladus
Haqicria
Onychoscpalum
Cafacolea
Colcocarva
Lcnidobolus
Kulima
Empodisma
A lexgcorgea
vpoiacna
Dapsilanthus
Apodasmia
Leptocamus
Chactanthus
Meeboldina
S ten o tails
Taraxis
Witrifrcdia
Stabcroha
Ischvrolepis
Thamnocnortus
Rhodocoma
Elegia
Chondropetalum
Dovca
Askidiosperma
Platvcaufos
Restio
Calopsis
Ceratocarvum
Willdenowia
Hypodiscus
Cannomois
Nevillea
Hvdrophilus
A'nthochortus
Mastersiella
CaJorophus
Sporadanthus
Lyginia < -
Loxocarya clade
Centrolepidaceae
Desmocladus clade
Leptocarpus clade
African clade
consensu s tree from parsimony analysis of morphological data for genera. Joinvillea
tMmrn formS the out 8 rou P- Arrows mark Lyginia and Hopkinsia. (Modified from Linder
et al. ZUUu).
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
481
Table 1. Morphological and phytochemical characters of Hopkinsia, Lyginia, Anarthria and
Restionaceae.
+ presence of character; - absence; * considered to be an apomorphic condition; ** autapomorphy,
i.e. sole occurrence among these taxa (some characters are marked as autapomorphies although common
to other taxa, e.g. 2-sporangiate anthers shared by Restionaceae and Centrolepidaceae). Character states
are defined so that conditions considered to be apomorphic are scored V. Several characters of related
organs are listed together where these are fully correlated among these taxa. Rare conditions are in
parentheses.
Character
dioecy
root hairs not persistent and lignified
culm sclerenchyma cylinder and
parenchyma sheath
leaf lamina reduced
leaves equitant
oblique epidermal cells and stomates
'false pillar cells' in chlorenchyma,
not around substomatal cavities
flowering culms not branched
inflorescence with caducous spathe
hyaline perianth and bracts
stamen filaments fused
anthers 2-sporangiate
pollen grain without graminoid annulus
ovary unilocular
stigmatic branches further branched
fleshy pedicel; drupaceous fruit
seed with spinules and median flange
mitotic metaphase chromosome length
(length of individual chromosomes)
starch in megagametophyte
fructan oligosaccharides in rhizome
silica absent from culm
rhombic crystals present
allose oligosaccharides present
flavones present
Hopkinsia
+*
+*
+
*
+ *
*
1.7-2.7 pm
?
+
Anarthria Restionaceae
+* +*/(-)
+/-
+*
+* - +*
_ | -k _
* _ _
_j_* *
+*/-
+*/-
*
+ */-
Lyginia
+*
+*
| -k * _ _
3-4 pm 3-7 pm 0.7-2 pm
+ - +
_i_* ★ _
+ +
+* +*
★ * _ _
+ +/-
(2) Exclusion from Anarthriaceae
Anarthria was separated from Restionaceae (Cutler & Shaw 1965) largely on the basis
of its fundamentally different culm anatomy. It lacks a sclerenchyma cylinder and
parenchymatous sheath isolating the chlorenchyma from the pith, having separate
sclerenchyma masses that surround each vascular bundle. The leaves are not reduced
but are highly distinctive in being equitant, with one margin representing the keel of
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a folded and fused structure. Hopkinsia and Lyginia differ strikingly from Anarthria in
these features of vegetative morphology and anatomy. The culms have both a
sclerenchyma cylinder and parenchyma sheath. Tire leaves are not equitant; there is a
much reduced dorsiventral lamina, often in the form of a mucro or awn at the apex of
the sheath. The leaf laminae of juvenile plants or those resprouting after fire are often
less reduced, but the lamina always has dorsiventral structure. The inflorescences also
lack a large enclosing spathe, as shown by Anarthria. All three genera possess sand¬
binding roots with long, persistent and lignified root hairs, but Pate and Delfs (1999)
indicate that this character may be plesiomorphic and loss of this epidermis in many
Restionaceae is considered to be apomorphic.
Although the analyses of DNA data indicate a close relationship between Anarthria,
Hopkinsia and Lyginia, if these genera lack morphological synapomorphies they fail to
form a family with recognisable apomorphic characteristics. The paucity of
synapomorphies is shown by the tree based on morphological data (Fig. 2), in which
the three genera are placed ungrouped at the base of the tree. Several of the features
that these genera share (Table 1) appear to be plesiomorphic in the Poales; these
include the 2-lobed, 4-sporangiate anthers and pollen with graminoid-type aperture.
The 23 characters (or character groups) listed in Table 1 reveal only the absence of silica
in the culms as a probable synapomorphy supporting the group ( Anarthria, Hopkinsia,
Lyginia) in contrast to Restionaceae.
(3) One new family or two?
Within the Anarthria clade, DNA data show ( Hopkinsia, Lyginia) as sister to Anarthria
(Fig. 1) but, as noted above, they fail to group in trees based on analysis of
morphological features (Fig. 2). Hopkinsia and Lyginia present very dissimilar
morphological features, and many of the features they share are plesiomorphic. They
share the synapomorphies of culm sclerenchyma cylinder and parenchyma sheath
(homoplasious with conditions in Restionaceae), but these features do not
convincingly associate them as a family in the face of many notable autapomorphies
(Table 1).
Distinctive autapomorphies for Hopkinsia include the thickened pedicel, long stigmatic
branches that are themselves branched, and fruit with both fleshy and woody layers.
Other features that are paralleled in some Restionaceae but are autapomorphic within
the Anarthria clade are the hyaline perianth and bracts, reduced gynoecium with a
single loculus and style, and indehiscent fruit. Linder (1992) observed that the ovary
of Hopkinsia differs in structure from other unilocular ovaries in Restionaceae.
Lyginia is marked by epidermal cells and stomates that are oblique to the culm surface,
chlorenchyma of alternating blocks of thin-walled photosynthetic cells and thick-
walled non-photosynthetic cells ('false pillar cells', not bordering substomatal cavities
and not homologous with Restionaceous pillar cells), fused staminal filaments, unique
seed ornamentation of cuticular spinules and a cuticular flange encircling the seed in
the median vertical plane.
We conclude, from the molecular and morphological evidence now available, that
Hopkinsia, and Lyginia cannot satisfactorily be included in either Restionaceae or
Anarthriaceae but that they do not form a single, well defined, new family. The
families Hopkinsiaceae and Lyginiaceae are therefore here established. The situation
is comparable to the recent recognition of two other new monocot families,
Anemarrhenaceae and Behniaceae (Conran et al. 1997), based in part on relationships
(oi lack of close relationship) suggested by molecular sequence data. Since the
Hopkinsiaceae and Lyginiaceae were not recognised by Kubitzki (1998), information
is provided here covering aspects treated for other monocot families in the Families and
Genera of Flowering Plants vol. 4.
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
483
Key to families of Poales
Restionaceae and allied families are generally included in the order Poales, but the
circumscription of that order has varied in different classifications. The Angiosperm
Phylogeny Group (APG 1998) included 15 families in Poales. The following key,
however, covers only the more restricted circumscription of seven families by
Dahlgren et al. (1985), together with the two new families now described. This range
includes the families considered to be closest relatives of the Restionaceae and of the
members of the Amrthrin clade.
1 Anthers 2-sporangiate, not lobed, with 1 dehiscence slit . 2
1* Anthers 4-sporangiate, 2-lobed, opening by 2 longitudinal slits . 3
2 Flowers mostly with tepals; male flowers with 2-3 stamens, females with 1-3 carpels;
pollen endexine present . Restionaceae
2* Flowers naked; males consisting of a single stamen; females of one carpel; pollen
endexine lacking. Centrolepidaceae
3 Fruit a drupe. 4
3* Fruit a capsule, nutlet, achene or follicle. 6
4 Leaf sheaths closed; leaf blades larger than sheaths and ending in a tendril .
. Flagellariaceae
4* Leaf sheaths open; leaf blades larger or smaller than sheaths and not ending in a
tendril . 5
5 Stems terete, without scabrid margins, unbranched below the inflorescence; leaf
blades plicate, larger than sheaths; plants with hermaphrodite flowers; ovary
trilocular; stylar branches or sessile stigmatic areas 3; stigmas covered with short
papillae . joinvilleaceae
5* Stems flattened, with scabrid margins, repeatedly branched; leaf blades not folded,
smaller than sheaths; plants dioecious; ovary unilocular; style single; stigmas covered
with long stigmatic branches that are themselves branched . Hopkinsiaceae
6 Flowers bisexual, each with a 2-keeled bract (palea) opposite the subtending bract
(lemma). Tepals ('lodicules') in one whorl (3 or fewer) . Poaceae
6* Flowers unisexual, with 1 or more bracts; bracts not as above (not 2-keeled). Tepals
in two whorls (mostly 3 + 3 or 2 + 2). 7
7 Monoecious with male and female flowers together in the same spikelet; stamens 4
or 6; culms striate with deep narrow crypts between broad sclerenchyma-capped
ridges.*. Ecdeiocoleaceae
7* Dioecious; stamens 3; culms not striate or with striations formed by narrow ridges
of sclerenchyma not by deep crypts .. 8
8 Leaf blades elongated, equitant (as if folded longitudinally with the 2 inner surfaces
fused, so that the abaxial margin represents the leaf keel and the blade is vertically
orientated); culms striate; stamen filaments not fused into a column .
. Anarthriaceae
8* Leaf blades of adult plants reduced to an awn at the apex of the sheathing base, not
equitant; culms not striate; stamen filaments fused into a column . Lyginiaceae
484
Telopea 8(4): 2000
Description of Hopkinsiaceae and revision of Hopkinsia
Hopkinsiaceae B.C. Briggs & L.A.S. Johnson, fam. nov.
Herbae dioeciae, perennes, rhizomatosae; culmi compressi, marginibus scabris; folia
dorsiventralia, reducta, vaginiformia; flores tepalis 6, scariosis; flores masculini
staminibus 3; antherae bilobatae, dorsifixae, sporangiis 4, utroque lobo per rimam
longitudinalem dehiscentes; filamenta antherarum discreta; grana pollinium aperturis
angustis; flores feminei ovario uniloculari et stylo unico; ovulum unicum; fructus
parva drupa.
Type and only genus: Hopkinsia W. Fitzg.
Perennial, glabrous, dioecious herbs with sympodial growth of rhizomes and culms;
caespitose with short rhizomes and forming large dense tussocks, or with long
rhizomes and forming diffuse patches of widely separated culms. Rhizomes covered
by appressed scales. Culms branched, smooth and terete near the base, becoming
flattened and concavo-convex above, with scabrous lateral margins. Leaves not
caducous, consisting of a sheath with a linear reduced lamina; sheath split to the base,
scarious, margins membranous; auricles scarious or membranous. Inflorescence
bracts: lower ones similar to culm sheaths but smaller, the uppermost hyaline. Flowers
wind-pollinated, not in spikelets, each in the axil of a membranous bract, not
dorsiventrally or laterally compressed; perianth of 6 membranous tepals in two
whorls. Male flowers: stamens 3, opposite the inner tepals; filaments free; anthers
dorsifixed, exserted, apically notched, 2-lobed, 4-thecate, latrorse, each lobe apiculate
and dehiscing by a longitudinal slit; the lobes attached below the midpoint to the
connective which is c. V 5 as long as anther; spirally twisted after dehiscence; pistillode
absent. Female flowers lacking staminodes; ovary superior, shortly stipitate, 1-locular,
not angled; the style single, stout, densely covered adaxially (except for a narrow
adaxial groove, i.e. stigma double-crested) by long stigmatic branches that are
branched and papillose, style attachment slightly excentric; the ovule solitary,
pendulous, orthotropous, bitegmic and tenuinucellate. Fruit a globose or elliptic small
drupe with a thin fleshy outer pericarp and a hard inner pericarp (although indehiscent
the slightly thickened and pale carpel margins are discernible and readily separable on
dissection); shed with a short fleshy pedicel and bracts and perianth attached.
Vegetative structure and anatomy: the culms are the main photosynthetic organs of
the plants and resemble those of Restionaceae in general structure (Cutler 1969;
Gilg-Benedict 1930; Cheadle & Kosaki 1975; Pate & Meney 1999; Meney, Pate &
Hickman 1999). The culm epidermis is tanniniferous and heavily thickened on the
outer wall; stomates are sunken, overarched by cuticular projections of epidermal
cells; substomata 1 cavities extend deeply into the chlorenchyma. The chlorenchyma
consists of 3-4 layers of short palisade cells with little or no development of pegs;
pillar cells, protective cells and sclerenchyma girders are absent. To the inside of the
chlorenchyma is a parenchyma sheath, l(-2) cells thick, surrounding the strongly
developed sclerenchyma cylinder. The outer vascular bundles are small and
embedded in the sclerenchyma, the inner bundles are larger with narrow
sclerenchyma sheaths. The central tissue is parenchymatous with patches of thin-
walled cells but no central cavity. Silica is apparently absent. The roots are sand¬
binding with a dense covering of persistent root hairs. The anatomy of rhizomes and
roots is illustrated by Meney, Pate and Hickman (1999).
Inflorescence and flower structure: inflorescences are (in the terminology of Briggs &
Johnson 1979) blastotelic (not terminated by a flower) and anauxotelic (axes not
growing on after flowering) as described for Restionaceae by Kircher (1986).
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
485
The arrangement of axes, especially after flowers have fallen, may suggest sympodial
growth but we interpret tire inflorescence structures as not sympodial. Flowers are not
in spikelets; flowers or flower-clusters are in the axils of small, ovate, acute, hyaline
bracts at two or several closely-spaced uppermost nodes of the culms and the lateral
branches. Male inflorescences bear clusters of several flowers on short slender axes;
female inflorescences are similar but bear a single flower in such a position. Both male
and female inflorescences are terminated by very small abortive axes (at the base of
the uppermost flower) bearing minute bracts. Male and female flowers are described
above for the family.
Pollen and embryology: the pollen grains are spheroidal to oblate, ulcerate, with a
polar diameter of 24-28(-33) pm. (Chanda 1966; Ladd 1977). The pollen walls show a
well-developed foot-layer, columellate interstitium, and a well developed tectum,
with numerous pores ('scrobiculi') penetrating the tectum and the foot-layer, as well
as other pores ('puncta') that penetrate the tectum only. Tire pollen surface shows
micro-verrucate ornamentation. The aperture is of graminoid type (Linder & Ferguson
1985), the annulus with a heavily thickened foot-layer, and the aperture diameter less
than 3-4 pm. A thin layer of endexine covers the ulcus and extends some distance
under the margins. The endothecium of the anthers contains spiral thickenings that
are distinctive in forming perfect helices that are doubled (Manning & Linder 1990).
The integuments are impregnated with tannins and the nucellar epidennis consists of
isodiametric cells (Kellogg & Linder 1995); the small, densely packed cortical cells of
the ovary resemble those of Lyginia (Linder 1992). The stigmas are dry. Embryo
development has not been studied.
Karyology: only a count of 2 n = 18 for H. adscendens, here published, is available.
(Fig. 3).
Fruit, seed and reproductive biology: flowers are wind-pollinated; the pollen is
released explosively and is normally completely expelled from the anther. The
dispersal unit is the fruit; a small drupe with thin fleshy pericarp, shed with a short
stout fleshy pedicel that contains oil and presumably functions as an elaeiosome. The
testa is fused to the pericarp, but with a white layer that may be equivalent to the seed
coat of related groups such as Lyginiaceae. Germination is hypogeal with the
I-1
Fig. 3. Hopkinsia adscendens, mitotic metaphase, 2 n = 18 (prepared as in Briggs 1966); voucher: Briggs
358. Scale bar = 5 pm.
486
Telopea 8(4): 2000
epicotylar rhizome developed (Pate & Meney pers. comm.; Pate & Meney 1999). The
rhizomatous habit facilitates resprouting after fire and, with the sand-binding roots,
serves to anchor these floodplain species which are subject to wash-aways of the
sandy substrate.
Phytochemistry: Hopkinsiaceae resemble some Restionaceae ( Lepyrodia and
Sporadanthus ) in having a simple flavonol pattern but lack flavones (Williams et al.
1997). Quercetin and isorhamnetin are reported in both Hopkinsia species;
proanthocyanidins and flavonoid sulphates are absent.
Distribution: one genus of two rare species on deep, seasonally moist, sandy soils,
that are oligotrophic or somewhat saline, in the southern half of Western Australia,
mostly near watercourses in regions of rather low rainfall.
Systematics and relationships: since the first description of species of Hopkinsia, this
has been included in Restionaceae. As mentioned above, DNA sequencing of the
chloroplast genes (Briggs et al. 2000) has now given convincing evidence that
Hopkinsia is more closely related to Anarthriaceae than to Restionaceae.
Hopkinsiaceae possess various features considered plesiomorphic within Poales: the
inflorescence with flowers not in spikelets, flowers with tepals not reduced in number,
tetrasporangiate anthers, graminoid pollen, presence of flavonoids but absence of
flavones. They differ from Restionaceae in the distinctive wiry, branching culms and
have a unique combination of features of culm anatomy (lacking pillars, girders and
protective cells; chlorenchyma of 3-4 layers of cells; silica absent). They share with
Lyginiaceae, and with various other Poalean families, many features considered
primitive in the Poales. Similarities to Lyginiaceae include the conspicuous sand¬
binding root-sheath of persistent, lignified, long root hairs (Pate & Meney 1999) and
the stout styles with stigmatic branches (Linder 1992), but they differ markedly in
habit, vegetative and floral morphology, fruits and seeds. They have none of the highly
distinctive features of the leaves and culm anatomy of Anarthriaceae or of
Ecdeiocoleaceae.
Hopkinsia
Hopkinsia W. Fitzg Western Australia Nat. Hist. Soc. 1: 33 (1904).
Type species: H. anoectocolea (F. Muell.) D.F. Cutler
Description and distribution: as for the family.
Key to species
1 Caespitose, forming large many-stemmed tussocks; rhizomes stout, 5-9 mm diam.;
culms 35-100 cm tall, densely crowded on the rhizomes; culm internodes (3—)6—11 cm
long; culm sheaths 2.5-3.5 cm long; 2-4 inflorescence branches arising at each upper
no ^ e .,.. 1. H. anoectocolea
1 Not caespitose; culms few and widely separated, arising from elongated horizontal
or ascending rhizomes; rhizomes slender, 1.2-2.5 mm diam.; culms 15-50 cm tall; culm
internodes 1.5-7 cm long; culm sheaths 0.7-1.5 cm long; inflorescence branches
arising singly at upper nodes . 2. H. adscendens
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
487
1. Hopkinsia anoectocolea (F. Muell.) D.F. Cutler, Kew Bull. 21: 61 (1967).
Basionym: Lepyrodia anoectocolea F. Muell., Fragm. 8: 78 (1873).
Type: Western Australia: W. A., /. Dr [Drummond] 6 (lecto, here selected MEL 14642; iso
MEL 14640-1, 14643, 15097, B, BM, K, P). [Some isotypes may be $: such plants are
difficult to distinguish from 6 when fruits have fallen.]
Anarthria calovaginata Gilg in Engl., Bot. Jahrb. Syst. 35: 87 (April 1904). Hopkinsia
calovaginata (Gilg) Gilg-Ben., Nat. Pflanzenfam. 2,15a: 17 (1930).
Type: in distr. Avon pr. Tammin ... Dliels] 5090 5 , 25 Oct 1901 (iso PERTH).
Hopkinsia scahrida W. Fitzg., J. Western Australia Nat. Hist. Soc. 1: 35 (May 1904).
Type: Cunderdin, saline spots, IV. V. Fitzgerald, Nov 1903 6 9 (syn PERTH 2067137,
1050109; isosyn B, NSW 406892, 413034).
Caespitose, forming large tussocks, sometimes to 1 m diam. Rhizomes horizontal, up
to 30 cm long or more, 5-9 mm diam., nodes 3-8 mm apart, bearing crowded culms
with their bases contiguous; covered with short glossy brown scales 5-8 mm long.
Culms 35-100 cm tall, 0.5-2 mm diam.; internodes (3-)6-ll cm long; young culms
pink, turning blue-green; old culms grey-green, glaucous, rigid and wiry; branching
except at the lowest nodes, with (1 —)2—4 lateral branches at each node; base with
several erect scales 1.5-6 cm long. Culm sheaths lax, sometimes recurved, reddish
brown, oblong, broadest near the middle, 2.5-3.5 cm long; auricles scarious; lamina
2-15 mm long. Inflorescence branches 2-4 at each upper culm node, flowers shortly
pedicellate, in clusters of 4—18 toward the apex of branches, subtended by broad-
deltoid, acute, hyaline bracts; outer bract c. 3 mm long, inner bract c. 1.5 mm long.
Male flowers: tepals ovate, concave, reddish or yellowish brown, soft, hyaline, acute;
outer tepals 1.3-1.9 mm long; inner tepals 2-2.6 mm long. Female flowers: tepals
similar to males, ± equal, 2.5-3 mm long. Fruit 2.2-2.7 mm long, 1.8-2.5 mm wide, red
when young, aging to brown, on a stout pedicel; seed surface patterned with convex
isodiametric angular cells.
Distribution: occurs in Western Australia north of Perth, at and near the Arrowsmith
River and eastwards at sites from the Mortlock River to Mekering and Tammin. In
flood-beds of watercourses, on deep sands or saline and clayey sands. Regenerates
after fire or flood by resprouting.
Conservation status: vulnerable (CALM code P2), with widely separated small
populations, sometimes locally abundant but occurring in habitats of limited
occurrence in regions of much clearing of natural vegetation and salinisation (Meney,
Pate, Dixon, Briggs, & Johnson 1999).
Selected specimens examined: Western Australia: Irwin: Arrowsmith Lake, George 12931, 9 Dec
1974 6, 9 (PERTH); Drummonds crossing. Brand Hwy at Arrowsmith River, Briggs 6406 6,6410
9, 27 Sep 1976 (NSW, AD, CANB, K, MEL, PERTH), Briggs 8618, Johnson, Meney, Linder & Pate, 8
Sep 1990 6 (NSW, BOL); 36 km NNW of Eneabba, Briggs 9366 6,9367 9 & Pate, 6 Oct 1995 (NSW,
NBG, PERTH). Avon: 3 km E of Meckering, Wilson 6413, 11 Nov 1968 9 (PERTH, K, NSW); Waeel,
Mortlock River, Wilson 6395, 23 Nov 1967 6 (PERTH, K, NSW); c. 18 km WNW of Cunderdin, Haegi
1164, 3 Oct 1976 (PERTH); 6.5 miles [11 km[ E of Cunderdin, 0.5 miles [1 km] E of 103 mile peg,
Briggs 202 6,203 9,5 Sep 1966 (NSW, CANB, K, MEL, PERTH); 86-87 mile peg, E of Meckering,
Gardner, 14 Dec 1945 6, 9 (PERTH).
2. Hopkinsia adscendens B.G. Briggs & L.A.S. Johnson, sp. nov.
A H. anoectocolea combinatione characterum sequentium distinguitur: rhizomata
angusta; culmi 15-50 cm longi, distantes vel aliquot aggregati; intemodia brevia, 1.5-7 cm
longa; vaginae culmorum breves, 0.7-1.5 cm longae; ramuli inflorescentiorum
singulatim in quoque nodo superiore.
488
Telopea 8(4): 2000
Type: Western Australia: Oldfield R., 61.7 km E of Ravensthorpe, 0.5 km N of bridge
on Hassell Hwy, B. Briggs 9343, J. Pate & K. Meney, 3 Oct 1995 9 (holo NSW; iso MO,
NBG, PERTH).
Forming diffuse patches of widely separated culms arising singly or several together
from long slender horizontal or ascending rhizomes which are up to at least 30 cm
long, 1.2-2.5 mm diam, internodes 1-3 cm long. Culms 15-50 cm tall, 0.8-1.5 mm
diam.; intemodes 1.5-7 cm long; young culms pink, turning blue-green; old culms
grey-green, glaucous, rigid and wiry', branched from near base, a single lateral branch
per node. Sheaths erect, oblong-spathulate, broadest toward the apex, slightly lax,
scarious, red-brown, 0.7-1.5 cm long, truncate, auriculate; lamina 0.5-3 mm long.
Inflorescence branches singly at upper nodes; flowers in clusters of 2-4 at the apex of
inflorescence branches, subtended by broad-deltoid, hyaline bracts almost as long as
the flowers. Male and female flowers 1.5-3 mm long, outer tepals 1.6-2.0 mm long,
inner tepals 2.0-2.6 mm long. Fruit ovoid, c. 2.7 mm long. Chromosome number: 2 n = 18
(Briggs unpublished, voucher Briggs 358). (Fig. 4.)
The epithet is from the Latin adscendens, ascending, referring to the rhizomes.
Distribution: occurs in Western Australia in a few locations from Chillinup (NE of
Albany) to the Esperance district. Grows in heath on sand or peaty sand in the
floodplain of watercourses in a region of moderately' low rain fall, often with eucalypts,
Banksia and Nuytsia. The occurrence of male and female plants suggests that
individual clones may be as large as 200 m across. Regenerates after fire by
resprouting.
Conservation status: endangered (CALM code R), limited to an uncommon habitat
type and occurring in regions of much clearing of natural vegetation and salinisation
(Meney, Pate, Dixon, Briggs & Johnson 1999; Pate 2000). Listed as at some risk but
poorly known (code 3K) by Briggs and Leigh (1996), but this appears to understate the
threats to this species.
Specimens examined: Western Australia: Eyre: 9 km E of Chillinup Homestead, Nezvbey 4882, Oct
1975 (PERTH); Oldfield R. (details as for type), Briggs 9342 et al. 6 (NSW, MO, NBG); c. 18 km
NNW of Young River crossing on Ravensthorpe -Esperance main road, Jackson 1437, 16 Oct 1968
<3 (AD, NSW); Lort River, c. 65 km W of Esperance, Orchard 1472, 11 Oct 1968 S (AD, NSW,
PERTH); Lort R., Esperance -Ravensthorpe Rd, Briggs 426 ,11 Sep 1966 6 (NSW, PERTH), Meney
393, 5 Mar 1993 6 (NSW); 40 miles [67 km] W of Esperance on Ravensthorpe road, Briggs 420,
11 Sep 1966 $ (NSW, K, PERTH), 421 6 (NSW, PERTH); 18 km NNW of Young R. crossing on
Ravensthorpe-Esperance Rd, Jackson 1437 ,16 Oct 1968 6 (PERTH, K, NSW); c. 10 miles [16 km) E
of Esperance on Cape Le Grand road, c. 1.5 miles [3 km] W of Cape Le Grand turn-off, Briggs 358,
9 Sep 1966 6 (NSW, MEL), 358a (NSW).
Description of Lyginiaceae and revision of Lyginia
Lyginiaceae B.G. Briggs & L.A.S. Johnson, fam. nov.
Herbae dioeciae, perennes, rhizomatosae vel caespitosae; culmi teretes; folia
dorsiventralia, reducta, vaginiformi; flores tepalis 6; flores masculi staminibus 3;
antherae bilobatae, dorsifixae, sporangiis 4, utroque lobo per rimam longitudinalem
dehiscentes; filamenta antherarum connata; grana pollinium aperturis angustis; flores
feminei ovario triloculari; ovulum unicum in quoque loculo; stylus crassus, in dimidio
superiore 3-partitus; fructus capsularis, triloculatus, loculicidus; semina alba, spiculis
minutis et crista mediana ornata.
Type and only genus: Lyginia R. Br.
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
489
Fig. 4. Hopkinsia adscendens. a-d, male: a, flower bud with bracts; b, flower cluster; c, flower;
d, inflorescence (Briggs 421). e-g, female: e, habit; f, flower; g, dissected flower with bracts, 6 tepals
and gynoecium (Briggs 420). Scale bar: a, b, c, f, g = 4 mm; d, e = 6.7 cm.
490
Telopea 8(4): 2000
Perennial, rushlike, dioecious herbs with sympodial growth of rhizomes and culms,
glabrous apart from cilia often on margins of culm and rhizome sheaths, spathes and
glumes; caespitose with short or erect rhizomes and forming small or large, dense or
± open tussocks, or with long creeping rhizomes and forming diffuse patches of
widely separated culms. Rhizomes covered by appressed, overlapping scales. Culms
unbranched, terete, smooth or finely rugose, with several nodes. Leaves reduced to
appressed sheaths, split to the base, persistent, rigid, acuminate; with a rigid, usually
black, incurved awn representing the reduced, dorsiventral lamina. Flowers not in
spikelets, singly or on short branching lateral axes in the axils of spathes at one or
several, usually closely spaced, uppermost culm nodes; wind pollinated, shortly
pedicellate, subtended by a bract; spathes persistent, rigid, acute or acuminate, mostly
partially obscuring the flowers; perianth of 6 tepals in two whorls, glabrous or ciliate
at the tips. Male flowers subtended by a hyaline bract as long as the tepals;
dorsiventrally compressed; tepals hyaline, blunt or acute or acuminate, glabrous or
ciliate at apex; outer lateral tepals keeled and the other outer and three inner tepals ±
flat; stamens 3, opposite the inner tepals; filaments connate into a stout tapering solid
column; anthers tetrasporangiate, free, dorsifixed, exserted, extrorse, attached below
the midpoint to the top of the column, apically notched, 2-lobed, often twisted after
anthesis; connective c. '/ 3 anther length; each lobe apiculate and dehiscing by a
longitudinal slit; pistillode absent. Female flowers not dorsiventrally or laterally
compressed, shortly pedicellate; tepals rigid, acute or acuminate, not keeled, the
margins toward the base hyaline, inner tepals shorter than outer; staminodes absent;
ovary superior, 3-locular, 3-angled; style branches 3, fused in the lower third or half,
stout, densely covered (except for a narrow adaxial groove [i.e., stigma double-
crested] and an abaxial ridge) by stigmatic papillae that are papillose; stigmas dry;
ovules solitary in each loculus, placentation axile, pendulous from summit of loculus,
orthotropous, bitegmic, tenuinucellate. Fruit a woody, 3-locular, 3-angled, loculicidal
capsule, style base persistent as a beak. Seeds globose, flattened near the hilum,
encircled in the median vertical plane by a hyaline flange; the outer layer white,
readily detached; the surface with fine short spines or hooks arising from the corners
of minute concavities.
Vegetative structure and anatomy: Lyginiaceae are evergreen, rush-like plants but the
species differ in growth habit and regenerative response. Two are caespitose but one
has extensive rhizomes producing patches of widely spaced culms; one is an obligate
seeder but two are resprouters (Bell & Pate 1993; Meney, Pate & Hickman 1999).
Growth is sympodial, the rhizomes covered by dark brown, usually glossy scales. The
sheaths surrounding the base of the culms develop a pattern of transverse cracks. The
culms are the main photosynthetic organs of the plants and are lime-green, yellowing
with age. The leaves are reduced to sheaths, which are persistent and conspicuously
or minutely ciliate or glabrous; with a stout tapering awn representing the reduced
amina, or with some development of the lamina in juvenile plants and in regrowth
a ter file. The roots are stout and sand-binding, and may attain great length with a
ense covering of persistent root-hairs; penetration to 4 m depth is reported by Pate
anc Meney (1999). The cilia on sheaths, spathes, bracts and tepals are stout, tapering,
multicellular, multiseriate, shortly papillate but unbranched.
The culms of the Lyginiaceae resemble those of Restionaceae in their specialisation for
tie combined functions of mechanical strength, photosynthesis, and resistance to
mm a ^u dUe t0 desiccation ( Cutler 19 69; Gilg 1891; Cilg-Benedict 1930; Malmanche
eac ^ e f955; Cheadle & Kosakai 1975). The stomates are oblique, sunken,
overarched by the radially-elongated epidermal cells. Under the epidermis is the
ayered chlorenchyma of palisade-like cells, with peg-cells in the substomatal
regions, interspersed with areas of thick-walled non-photosynthetic cells (not
or ermg substomatal cavities); pillar and protective cells of the types found in
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
491
various Restionaceae and sclerenchyma girders are all absent. (Cutler [1969 Fig 351
identifies thick-walled cells in the chlorenchyma of Lyginia as pillar cells, but notes [p
234] that They are derived from palisade chlorenchyma cells [rather than cells of the
parenchyma sheath] and therefore cannot be regarded as true pillar cells.') To the
inside of the chlorenchyma is the parenchyma sheath, l(-2) cells thick surrounding the
sclerenchyma cylinder. The outer vascular bundles are embedded in the sclerenchyma
cylinder, the inner bundles are scattered in the pith. Some pith cells include slab¬
shaped crystals; there is no central cavity. The central ground tissue is
sclerenchymatous around the lignified bundle sheaths, and there are numerous
intercellular spaces. The anatomy of rhizomes and roots is illustrated by Meney, Pate
and Hickman (1999) and further description of morphology and anatomy given by
Pate and Meney (1999) and Pate and Delfs (1999).
Lyginia shows several distinctive anatomical features (Gilg 1891; Cutler 1969): the thin-
walled cells lining the substomatal cavity which are unlike those of genera in related
families that show differentiated substomatal cells, the oblique epidermal cells,
stomata at an angle to the epidermal cells, and slab-shaped crystals in some cells of the
central ground tissue. The absence of silica also contrasts with the usual condition in
Restionaceae. Cheadle (1955) considered the vessels of the shoot system (but not of the
roots) of Lyginia to show especially primitive characteristics.
Inflorescence and flower structure: inflorescences are blastotelic (not terminated by a
flower [Briggs & Johnson 1979]) and anauxotelic (axes not growing on after flowering);
with rigid, acuminate culm spathes at 1-several uppermost culm nodes, subtending
short flowering branches that may bear smaller spathes subtending very short
flowering branches; each flower subtended by a glume-like bract. Female
inflorescences are fewer-flowered and less branched than the males. Male and female
flowers are described above for the family.
Pollen morphology and embryology: the single pollen grains are spheroidal to oblate,
c. 30-38 pm polar diameter (Chanda 1966; Ladd 1977). The pollen walls show a foot-
layer, columellate interstitium, and a thin tectum, with numerous pores penetrating
the tectum and the foot-layer ('scrobiculi'), as well as pores ('puncta') that penetrate
the tectum only. The pollen surface has micro-verrucate ornamentation. The annulus
consists of a heavily thickened foot-layer, and the aperture is of 'graminoid' type
(Linder & Ferguson 1985) with a diameter about 7 pm. A thin layer of endexine covers
the ulcus and extends some distance under tine margins. The endothecium of the
anthers contains spiral thickenings but these often form U-shapes rather than
complete helices, showing similarity to the condition in Anarthriaceae (Manning &
Linder 1990). The ovary walls are impregnated with tannins. Linder (1992) observed
that the ovary is similar to that of Anarthria but has small densely packed cortical cells
more like those of Hopkinsia. The orthotropous, tenuinucellate ovules are bitegmic,
with the inner integument tanniniferous. Megagametophyte development is not
known beyond the 8-nucleate stage but there are three antipodals and large starch
grains in the mature megagametophyte (Rudali & Linder 1988). The development of
the embryo has not been studied.
Karyology: Lyginiaceae have the number of n = 6 (Johnson & Briggs 1981; Bell & Pate
1993) which is not known in the Restionaceae and the chromosomes are larger than in
the latter family, although not as large as in Anarthriaceae (Briggs 1966). [Johnson and
Briggs (1981) corrected a previous erroneous count by Briggs (1966)]. Bell and Pate
studied all species and observed that there 'were no apparent differences in size or
configuration of chromosomes among [them]'.
Fruit and seed: the capsule develops from a 3-locular ovary, with one seed per locule.
The seed has a readily detached, white outer layer. The surface is highly distinctive
492
Telopea 8(4): 2000
with cuticular spinules arising from the angles of palisade-like, very slender and
radially elongated cells of the outer layer of the testa; the cuticle is concave over the
outer surface of each cell. The flange encircling the seed in the median vertical plane
appears to be a projection of the cuticle and not a cellular structure.
Reproductive biology: the flowers are wind-pollinated and dioecious, with the seeds
serving as disseminules. The pollen is released explosively and normally completely
expelled from the anther. Flowering is autumn or spring (bimodal) (Meney & Pate
1999). Germination is hypogeal with the epicotylar rhizome developed (Pate & Meney
pers. comm.) Lyginiaceae occur in fire-prone environments, and the presence of an
obligate seeder species as well as resprouters, with distinctive patterns of resource
allocation, was reported and discussed by Pate and colleagues (Bell & Pate 1993;
Meney, Dixon & Pate 1999; Pate, Meney, Dixon, Bell & Hickman 1999).
Fungal associates: Lyginiaceae resemble Restionaceae in their resistance to
Phytophthora infection but a culm smut recorded as infecting all Lyginia species
(Websdane et al. 1994; Sieler et al. 1999) is sufficiently distinctive to have been
described as the new monotypic genus Websdanea (Vanky 1997). This smut has not
been recorded on other hosts. Meney et al. (1993) recorded the presence of mycorrhizal
associations for a brief period during winter in the roots of Lyginia ; mycorrhizae have
rarely been reported in Restionaceae.
Phytochemistry: Lyginia differs from members of related families in the presence of
the unusual sugar allose and of allosides, while resembling Restionaceae in producing
some simple flavonol glycosides (Williams et al. 1997). Quercetin and isorhamnetin
were reported in most samples; proanthocyanidins and flavonoid sulphates are
absent. Williams et al. also note that, as in some Restionaceae, 'the more chemically
advanced 8-hydroxy flavonol gossypetin ... is present in at least one species'. Unlike
Restionaceae, Lyginia produces significant amounts of fructan-type oligosaccharides in
its rhizomes and lacks starch (Pate, Meney & Dixon 1991).
Systematics and relationships: Lyginia has until now been included in Restionaceae.
As noted above, cladistic analysis of DNA sequence data from the chloroplast gene
) cL, the /n/L intron and the fnzL-F intergenic spacer (Briggs et al. 2000) do not
associate Hopkinsia and Lyginia with Restionaceae but show a robust clade of these two
genera with Anarthria. Within this clade Anarthria is sister to ( Hopkinsia , Lyginia).
As with Hopkinsia, the general similarity of Lyginia to Restionaceae in gross
morphology and anatomy, and its occurrence in a region of great diversity of
estionaceae, have led to its distinctiveness being overlooked. Lyginiaceae possess
various conditions that are considered plesiomorphic within Poales, in the
in oiescence, flowers (unreduced tepals, tetrasporangiate anthers, gynoecium), pollen
morp o ogy, ovule and flavonoids. However, they show distinctive apomorphies in
1 oresc fnce structure, connate staminal filaments, stomatal position, structure of the
chlorenchyma, and seed surface.
Distribution and habitats: the single genus Lyginia is widespread on well drained or
seasona y moist oligotrophic soils in the southern half of Western Australia, in regions
o mo erate to low rainfall. One species is rare but two are wide-ranging and common.
Lyginia
Lyginia R. Br., Prodr.: 248 (1810).
Lectotype. L. barbata R. Br. (typ. cons.) (In 1910, Lyginia R. Br. was conserved against
c toeno urn Labill., following a proposal by Janchen 1908), but this was subsequently
recognised as a superfluous conservation (Rickett & Stafleu 1959, p. 231). Schoenodum
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
493
tenax Labill. was based on specimens of two species. Brown selected the male
specimen as lectotype, basing Leptocarpus tenax (Labill.) R. Br. on it, while referring the
female syntype to Lyginia imberbis R. Br. Lyginia is no longer included among the
nomina conservanda genericorum, although it retains a conserved type.)
Description and distribution: as for the family.
Species within Lyginia have proved difficult to delimit, so that the names L. barbata and
L. imberbis have been defined variously, or the whole genus considered to be only a
single species. A major clarification came with the studies of Bell and Pate (1993) who
investigated the reproduction and regeneration of plants and distinguished five
morphotypes. These were characterised by differences in rhizome morphology, culm
height and density, spikelet morphology, response to fire, timing of first reproduction
in juveniles, and seed/ovule numbers and ratios. The species now recognised equate
to these units, as characterised by Bell and Pate, as follows:
L. excelsa sp. nov. = morphotype S: 'tall-culmed, densely tufted fire-sensitive
obligate seeder'.
L. imberbis R. Br. = morphotypes R] 'tall densely tufted fire-resistant resprouter'; R 2 :
'short flexuose-culmed, tufted resprouter'; and R 3 : 'weakly clonal, semi-tufted
resprouter'.
L. barbata R. Br. = morphotype Ry 'short-culmed, widely spreading clonal form'.
We have concluded that there is no basis for specific distinctions between
morphotypes R,, R 2 and R 3 . L. imberbis, as now recognised, is by far the most variable
and abundant of the species. It includes considerable local variation; for example,
plants from different sites often show consistent differences in culm height and
sinuosity, corresponding to three of the morphotypes distinguished by Bell and Pate
(1993). L. barbata shows an almost equally wide geographic distribution but is
restricted to a much narrower habitat range, whereas L. excelsa is known from only a
very few sites. Bell and Pate found differences between the morphotypes now
included in L. imberbis in seed/ovule ratios and resource allocation to reproduction,
but by far the largest differences in these variables correspond to the specific
distinctions now recognised. They noted the clear demarcation between the groups
now specifically distinguished, regarding Rj, R 2 and R 3 as together forming 'an
intermediate grouping of resprouter forms', distinct from the obligate seeder and
extensive clonal types.
The species occur sympatrically or in close proximity without evidence of
intergradation (Bell & Pate 1993). L. excelsa occurs with L. barbata at Cataby. L. imberbis
and L. barbata are found together at various localities but are more often parapatric
since L. barbata is generally on more sandy and better drained sites. There is little
difficulty in identifying most collections that adequately sample the base of the plant.
Especially in the Eyre region, however, some plants are difficult to determine since L.
imberbis shows considerable variation in the denseness of the tussocks, the extent to
which culms are supported by short but distinct rhizomes, and in the development of
cilia. Elongated rhizomes are a reliable characteristic of L. barbata, although these do
not develop until about the fifth year of growth from seed (Bell & Pate 1993). Since
establishment from seed is infrequent in this strongly clonal species, juvenile plants
that would be difficult to identify are correspondingly rarely encountered (Pate 2000).
494
Telopea 8 ( 4 ): 2000
Key to species
1 Plants not caespitose, forming patches of widely spaced culms or small groups of
culms connected by long rhizomes; culm sheaths and inflorescence spathes ciliate
with stiff hairs. L. barbata
1* Plants caespitose, rhizomes erect or shortly horizontal, with crowded culms; culm
sheaths and inflorescence spathes glabrous or sparsely or strongly ciliate . 2
2 Culms 30-100 cm tall, straight or sinuous, intemodes 5-11 cm long; culm sheaths
and inflorescence spathes glabrous or sparsely ciliate with hairs 0.5-2 mm long;
bracts at base of culms closely appressed and with an awn-like lamina 2-10 mm
long; female inflorescence flowering at 1-5 upper culm nodes; fertile spathes each
subtending a single flower . L. imberbis
2* Culms 50-150 cm tall, straight, internodes 14-25 cm long ; culm sheaths and
inflorescence spathes ciliate with stiff hairs 3-6 mm long; bracts at base of culms ±
flat in the upper half and not closely appressed, with a small mucro or an awn 1-4
mm long; female inflorescence flowering at 4- 7 upper culm nodes; fertile female
spathes each subtending 1-3 flowers. L. excelsa
Lyginia barbata R. Br., Prodr. 248 (1810)
Lyginia barbata var. barbata
Type citation: (M) v.v.
Type: Western Australia: King George III d S d [Sound], R. Brown (Bennett No. 5838),
1802-5 5 , 9 (holo BM [annotated by Brown 'Nov genus inter Restionem']; iso BM, E, P).
[Note on typification. At the International Botanical Congress in 1999 a change was
accepted in Article 8.1 of the International Code of Botanical Nomenclature to avoid
unnecessary lectotype designations" narrowing down the type to a single individual
or fragment thereof' (Greuter & Hawkesworth 1999). The definition of a type (Barrie
& Greuter 1999) now emphasises that 'a specimen is a gathering, or part of a gathering,
of a single species ... or of multiple small plants'. Discussion at the meeting confirmed
the acceptability of male and female plants of the same taxon within a gathering for
purposes of typification. Lectotypes are here designated in some instances where what
appears to be a single gathering is mounted on more than one sheet and these are not
cross-referenced, but no selection of a lectotype is made between 6 and 9 specimens
of the same taxon mounted on a single sheet.]
Rhizomatous, forming large patches of widely separated culms or of groups of several
culms, 2-14 cm apart. Rhizomes, horizontal or ascending; 3-6 mm diam. including the
covering of dark brown, appressed, usually glossy scales; intemodes 0.5-1.3 mm long.
Culms straight and erect or sinuous, smooth or minutely rugose, 30-80 cm tall, 1.3-2
mm diam.; with 4-8 intemodes, each 5-18 cm long. Sheaths at base of culms
± truncate, 10-18 mm long, closely enwrapping the culms, ciliate, with an awn 4-8 mm
long. Culm sheaths 6-15 mm long, green or red-brown, truncate or tapering abruptly
at the apex, ciliate at apex with hairs 2.5-6 mm long, lax toward apex, with a curved,
lack awn 6—13 mm long. Culm spathes acute, rigid; 6-13 in male inflorescences, 1-4
in female inflorescences, each subtending only a single flower, or culms with several
closely spaced spathes but only the uppermost subtending a flower; spathes 4-12 mm
ong, ciliate with stiff hairs, with a rigid curved awn as long or longer than the spathe.
Lateral branches of male inflorescences condensed or extending beyond the culm
spathe and with small spathes conspicuous on the lateral branches, to 2.5 cm long,
ale flowers: tepals hyaline, narrow lanceolate, shortly ciliate at apex, acuminate or
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
495
Fig. 5. Lyginin. a-c, L. Imrbntn. a, habit; b, part of female inflorescence (Briggs 6765);
c, male inflorescence (Briggs 151). d-i, L. imberbis. d, male inflorescence (Krnuss 144 & Hoivitt);
e, f, male flower and dehisced anther (Coveny 8149); g, female flower with bracts (Coveny 8123);
h, capsule with bracts and tepals; i, dehisced capsule (Krauss 140 & Hoivitt). Scale bar: a = 7 cm;
b, c, d, g, h, i = 7 mm; e = 3 mm; f = 1.75 mm.
496
Telopea 8(4): 2000
acute or blunt, 4-6 mm long; filament column 5-7 mm long; anthers 2.5-4 mm long.
Female flowers: pedicel c. 1.5 mm long; tepals rigid, 4-6 mm long, shortly ciliate
toward apex; outer tepals ovate, with a slender black awn 3-5 mm long; inner tepals
narrow deltoid, acute; style 12-15 mm long. Capsule depressed globular, 3-angled, 5-6
mm long, 6-8 mm diam. Seeds c. 2 mm diam. Chromosome number 2n = 12 (Bell &
Pate 1993). (Fig. 5a-c).
Distribution: widespread on oligotrophic soils in the southern half of Western
Australia from Eneabba to Israelite Bay, on deep sands in well drained or seasonally
damp woodlands and heaths, often with Banksia shrubs. Commonly in better drained
habitats than L. imberbis. Regenerates after fire by resprouting. Clones may be long
lived and extend over areas 50-100 m across (Bell & Pate 1993).
Conservation status: widespread and common.
Selected specimens examined: Western Australia: Irwin: 25 km SSE of Eneabba, Briggs 9007, 8 Oct
1992 3 (NSW); 1 km W of Brand Hwy along Greenhead Road, Wilson 2688, 2 Oct 1979 9 (NSW,
PERTH); 5 km S of Badgingarra, Keighery 2563, 10 Sep 1979 $ (PERTH); Cataby, Beil & Pate s.n.,
Oct 1991 3, 2 (KPBG). Darling: c. 29 km NNW of Gingin, Briggs 6765, 11 Oct 1976 9 (NSW);
Melaleuca Park, c. 8 km WSW of Bullsbrook, Muir 830, 3 Sep 1977 $ (PERTH); Bayswater, Gardner,
Oct 1940 ? (PERTH); Cannington Swamp, Eicbler 15718, 27 Aug 1959 3 (AD, PERTH); Cannington,
b'j88 s 151, 3 Sep 1966 o’ (NSW); W of Yarloop, 3.9 km E of Old Coast Road on Johnsons Road, Hill
4594 6, 4595-6 9,10 Oct 1993 (NSW); 2.4 km SSW of Paynedale along Goodwood Rd, c. 14 km
SW of Donnybrook, Briggs 8310 & Johnson, 25 Oct 1988 9 (NSW); Ambergate, Busselton district,
Royce 2869, 18 Oct 1948 2 (PERTH); Bow River, S.W. Jackson, Jan 1913 3, 9 (PERTH); 9 km S of
Mt Barker, Blake 20879, 1 Sep 1959 3 (BRI, NSW); Denmark, Gardner s.n., Feb 1940 3 (PERTH);
Albany, Pieiss 1699, Oct 1840 3 (NSW). Eyre: below WestMt Barren, ESE of Ongerup, Coveny 3305,
10 Sep 1970 3 (NSW, CANB, K, NBG, PERTH); 0.8 km S of Gibson, Briggs 335, 9 Sep 1966 3 (NSW,
PERTH, K); Lucky Bay, E of Esperance, Briggs 367 3, 368 9 10 Sep 1966 (NSW, PERTH, RSA).
Lyginia imberbis R. Br„ Prodr. 248 (1810)
Lyginia barbata var. imberbis Mast, in A.P.C. de Candolle, Monogr. phan. 1: 303 (1878).
Type citation: (M) v.v.
Tcnn ^ estern Australia: King George III d S d [Sound], R. Brozvn (Bennett No. 5837),
1802-5 3, 9 (holo BM, annotated by Brown 'Restio'; iso BM, E, K, P).
Caespitose, forming small or large, dense or + open tussocks. Rhizomes short,
orizontal or ascending, with crowded culms, covered by dark brown ± glossy scales,
in' ton CreCt anC * stra '§* lt or sightly or strongly sinuous, smooth or minutely rugose,
cm tall, 0.8-1.4 mm diam.; with 4-6 internodes, each 5-11 cm long. Sheaths at
ase of culms 5-20 mm long, with a mucro or awn 1-5 mm long, glabrous or sparsely
a late with hairs c. 1 mm long, mostly closely enwrapping culms. Culm sheaths
... mm on S' green or red-brown, tapering sharply to the apex; glabrous or sparsely
a mte at apex with hairs 0.5-2 mm long, lax toward apex, with a curved black awn
. _ mm on S- Culm spathes acute, rigid, 1-11 in male inflorescences, 1-5 in female
in orescences, each subtending only a single flower, or culms with several closely
spaced spathes but only the uppermost subtending a flower; spathes 3-9 mm long,
glabrous or sparsely ciliate, with a rigid curved awn about as long as the spathe.
ateral brandies of male inflorescences condensed and not extending beyond the
spa e. a e flowers: tepals hyaline, oblong or narrow lanceolate, glabrous or sparsely
ciliate at apex, acuminate or acute or blunt, 5-7 mm long; filament column 7-8 mm
on & ant ers 2.5-3.5 mm long. Female flowers: pedicel c. 1.5 mm long; tepals rigid,
1 A T glabrous or sparsely ciliate toward apex; outer tepals ovate, with a
s ent er i ac awn 2.5-7 mm long; inner tepals deltoid, acute or acuminate; style 8-10 mm
°ng. apsu e depressed globular, 3-angled, 4-5 mm long, 5.5-7 mm diam. Seeds
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
497
c. 2 mm diam. Chromosome number 2 n = 12 (Johnson & Briggs 1981, voucher Gittins
1614b; Bell & Pate 1993). (Fig. 5d-i).
Distribution: widespread on oligotrophic soils in the southern half of Western
Australia from the Murchison River to Israelite Bay, in sand or peaty sand in
seasonally damp heath and woodland. Commonly in moister habitats than L. barbata.
Regenerates after fire by resprouting.
Conservation status: widespread and common.
In addition to the variable features noted by Bell and Pate (1993), plants at the northern
end of the geographic range (Murchison River and Kalbarri to just north of Eneabba)
show notably slender and straight culms; male plants mostly with numerous (7-11)
flowering spathes but females with few (1-2) spathes. Rather similar plants are found
elsewhere in the distribution, but the uniformity in this northern region is notable. At
Eneabba most plants are very different from those further north, having stout and
highly sinuose culms (type R 2 of Bell & Pate 1993), often with crowded spathes in 6
inflorescences.
Selected specimens examined: Western Australia: Irwin: Murchison House Station, Gittins 1614a
3, 1614b 9, Aug 1967 (NSW); 9 km ENE of Kalbarri, Krauss 140 & Hoivitt, 16 Apr 1989 9 (NSW,
MEL, PERTH); 31 km NNW of Eneabba, Krauss 144 & Howitt, 22 Apr 1989 3 (NSW); 37 km SE of
Kalbarri on Ajana road, Wilson 2666, 1 Oct 1979 3 (NSW, CANB, PERTH, RSA), 2665 9 (NSW,
CANB, PERTH); 3.7 km from Baline on road to Binnu, Briggs S881 & Johnson, 13 Aug 1991 9 (NSW);
11 km E of Dongara on Midlands Road, Briggs 7492 & Johnson, 30 Sep 1984 9 (NSW, CANB, MEL,
PERTH); Eneabba, Bell, Sep 19913, 9 (KPBG); Eneabba, Haegi 1928 & Powell, 1 Sep 1979 (NSW, AD,
CANB, K, PERTH, RSA). Darling: White Well Rd, 12 km due S of Ballidu, Briggs 9000a, 9000b 9,
7 Oct 1992 3, (NSW, PERTH); Kewdale, Covcny 8199, 7 Sep 1976 3 (NSW, K, PERTH), 8198 9
(NSW, BR1, CANB); Brook Rd, Wattle Grove, Covetty 8123, 6 Sep 1976 9 (NSW), 8149 3 (NSW);
South Street, Gosnells, Bell & Pate, Sep 1991 3, 9 (KPBG); W of Yarloop, 3.9 km E of Old Coast
Road on Johnsons Road, Hill 4597, 10 Oct 1993 9 (NSW); 2.4 km SSW of Paynedale along
Goodwood Rd, c. 14 km SW of Donnybrook, Briggs 8308 & Johnson, 25 Oct 1988 9 (NSW); Scott R.
Plains, junction of Cane Break Rd and Governor Broome Rd, Briggs 6506, 2 Oct 1976 3 (NSW).
Avon: 10.5 km E of Cunderdin, Briggs 2000, 5 Sep 1966 3 (NSW); Avon, Pritzel 894 3, 895 9, Nov
1901 (NSW, P); 30.6 km ESE of Pingelly, E side of Tutanning Reserve, Briggs 6655, 6 Oct 1976 9
(NSW). Eyre: Oldfield R., 62 km E of Ravensthorpe, Wilson 2820, 13 Oct 1979 3 (NSW, PERTH);
10 km WNW of Chillinup Pool, Newbey 4551a 3, 4551b 9, 29 Oct 1974 (PERTH, NSW); c. 9 km S
of Israelite Bay, just W of Lake Daringdella, Green 5104, 21 Nov 1986 9 (PERTH). Roe: c. 30 km W
of Mt Ragged, Wilson 2935,10 Sep 1964 3 (AD, NSW).
Lyginia excelsa B.G. Briggs & L.A.S. Johnson, sp. nov.
Herbae caespitosae; rhizomata haud producta; seminibus reproductio; vaginae basales
18-40 an longae; culmi 50-150 cm longi; vaginae culmorum 20-40 cm longae; vaginae
culmorum et spathae inflorescentiarum ciliatae, pilis 3-6 mm longis; spathae
plantarum feminearum 1-3-floribus.
Type: Western Australia: Brand Hwy N of Cataby, 0.2 km S of Mullering Road, B.G.
Briggs 9319, 30 Sep 1995 9 (holo NSW; iso CANB, K, PERTH).
Caespitose, forming large, dense tussocks to 30 cm across at base. Rhizomes short,
vertical or steeply ascending, bearing closely crowded culms and dark brown slightly
glossy scales. Culms erect, straight, 50-150 cm tall, 1-2.2 mm diam, smooth or
minutely rugose, with 3-5 intemodes each 14-25 cm long. Sheaths at base of culms,
5-20 mm long, ± flat in the upper half and not closely enwrapping the culms, with a
mucro 1-2 mm long or a short awn to 4 mm long. Culm sheaths 20-30 mm long, green
or brown, tapering gradually toward the apex, ciliate with stiff hairs 3-6 mm long, lax
toward apex, with a curved black awn 8-10 mm long. Culm spathes acute, rigid, 7-9
in male inflorescences, 4-7 in female inflorescences; each subtending a short, often
498
Telopea 8(4): 2000
branched, axis with 1-3 flowers, of which 2 may develop fruit; spathes 10-20 mm long,
ciliate with stiff hairs, with a rigid curved awn 6-14 mm long. Lateral branches of male
inflorescences generally extending beyond the spathe, to 15 mm long. Male flowers:
tepals hyaline, linear, sparsely ciliate at apex, acute, 6-7 mm long; filament column
c. 9 mm long; anthers c. 3.5 mm long. Female flowers: pedicel 1.5-2.5 mm long; tepals
rigid, deltoid, acute, sparsely ciliate toward apex; outer tepals (3-)5-8 mm long, with
a slender black awn c. 4 mm long; inner tepals 5-6 mm long; style 8-10 mm long.
Capsule depressed globular, 3-angled, 4—6 mm long, 6-8 mm diam. Seeds c. 2 mm
diam. Chromosome number In = 12 (Bell & Pate 1993). (Fig. 6).
The epithet is from the Latin excelsus, lofty or high, referring to the stature of the
plants, which are mostly taller than other species of Lyginia.
Distribution: occurs in Western Australia north of Perth, at a few sites near Cataby
and north of Badgingarra, on sand in dry heath and Banksia woodland.
Conservation status: vulnerable; known from only a few sites and in a region subject
to clearing of native vegetation. CALM code P. Obligate seeder species, made more
vulnerable since it is killed by even mild fires (Meney, Pate, Dixon, Briggs & Johnson
1999; Pate 2000).
Specimens examined: Western Australia: Irwin: 2.9 km N of Tootbardie Rd on Hwy 1, Briggs 8921
& Johnson, 15 Aug 1991 6 (NSW); junction of Brand Hwy and Cooljarloo Rd, N of Cataby, Meney
2191,22 Oct 1990 $ (progeny of this collection, raised from seed: NSW); Brand Hwy N of Cataby
(details as for type), Briggs 9317 8 (NSW), 9318 9 (NSW, PERTH), 9320 9 (NSW, K, PERTH);
Cataby, Bell & Pate, Oct 1992 9 (KPBG).
Excluded and illegitimate names
Lyginm tenax (Labill.) Steud. (as Lygynia), Nomencl. bot. ed. 1, 1; 501 (1821) =
Leptocarpus tenax (Labill.) R. Br.
Lyginia montevidensis Spreng. in Linn. Syst. veg. 4 (2): 26 (1827). Referred to 'species
ignotn ... excludenda ' by Masters in Candolle Monogr. phan. 1: 303 (1878).
Lyginia symphyonema F. Muell., Fragm. 8: 79 (1873), notn. illeg., based on an inclusive
concept of L. barbata R. Br., L. imberbis R. Br. and Schoenodum tenax Labill.
Acknowledgments
John Pate (University of Western Australia), Kathy Meney, Tina Bell, and Kingsley
ixon ( mgs Park and Botanic Garden, Perth) shared their knowledge with us most
generous y in discussions and in the field. The information they provided helped
greatly, especially in clarifying the classification of Lyginia. The DNA studies that
prov i ec part of the basis for distinguishing these new families also benefited from
p an s in tissue culture and seeds provided by Meney and Dixon. Thanks go to
co eagues w 10 gave valuable assistance, especially Adam Marchant, Simon Gilmore
3IU b ^ orter ( u 'h°se DNA sequencing studies were crucial to the conclusions
reac cc), legfried Kiauss and Barbara Wiecek. Horticultural colleagues, at the Mount
nnan otanic Garden and in Sydney, maintained relevant living collections. David
ac ay, icola Oram and Lesley Elkan were responsible for the illustrations and Peter
i son assisted with the Latin diagnoses. The opportunity to examine specimens on
oan or in otier herbaria, and extensive data provided by the Western Australian
ti anum ( RTH),contributed to the study. Grants from the Australian Research
ounci an ustralian Biological Resources Study supported part of the project,
omments on the manuscript from Peter Linder (University of Cape Town) and
mem ieis o the Telopea Editorial Committee improved the paper.
Briggs and Johnson, Hopkinsiaceae and Lyginiaceae
499
Fig. 6. Lyginia excelsa. a-d, female: a, inflorescence; b, culm sheath; c, flower; d, base of plant with
erect rhizomes, sheaths and culm bases (Briggs 9319); e, fruit {Briggs 9320). f-h, male: f, flower;
g, flower with tepals opened out; h, inflorescence {Briggs 9317). Scale bar: a = 1.5 cm; b, e = 1 cm;
c, f, g = 7.5 mm; d= 2.5 cm; h = 3 cm.
500
Telopea 8(4): 2000
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Manuscript received 31 January 2000
Manuscript accepted 15 May 2000
503
Systematic studies in the eucalypts. 10.
New tropical and subtropical eucalypts from
Australia and New Guinea
(Eucalyptus, Myrtaceae)
K.D. Hill and L.A.S. Johnson*
Abstract
Hill, K.D. and Johnson L.A.S., (National Herbarium of Neiu South Wales, Royal Botanic Gardens,
Sydney, Australia 2000) 2000. Systematic studies in the eucalypts. 10. New tropical and subtropical
eucalypts from Australia and Neiv Guinea (Eucalyptus, Myrtaceae). Telopea 8(4): 503-539. New
species described are Eucalyptus biterranea, E. macta, E. grisea, E. pantoleuca, E. glomericassis,
E. kenneallyi, E. costuligera, E. epruinata, E. limitaris, E. tephrodes and E. xerothermica. New
subspecies are recognised in E. leucophloia Brooker (subsp. euroa), E. oligantha Schauer (subsp.
modica), and E. pruinosa Schauer (subsp. tenuata). All new taxa fall within subgenus
Symphyomyrtus, in the three sections Transversaria, Exsertaria and Adnataria. New taxa are
mapped, and keys are presented in cases where existing keys will not discriminate taxa. Selected
taxa are illustrated.
Introduction
New taxa described here are tropical species from the three sections Transversaria,
Exsertaria and Adnataria of subgenus Symphyomyrtus of the genus Eucalyptus taken in
the sense of Hill and Johnson (1995), excluding the genus Corymbia as discussed
therein. The new taxa are from New Guinea and tropical regions of Queensland,
Northern Territory and Western Australia. Several of these taxa are treated as
undescribed species or subspecies in the semi-popular account of tropical taxa by
Brooker and Kleinig (1994). Our new taxa, discussed both here and by Brooker and
Kleinig were delineated by us during a comprehensive revisionary study of the
eucalypts, and were freely discussed with Ian Brooker in order to allow the treatment
in Brooker and Kleinig.
Terminology and nomenclature are as in previous papers in this series (see Hill &
Johnson 1995). Rare or threatened species are allocated conservation status codes
according to the system of Briggs and Leigh (1996). Species authorship is to be cited as
presented under each taxon described. They are not cases for the use of 'ex'. Keys are
presented in cases where existing keys will not discriminate taxa, either as
dichotomous keys or comparative tables. In other cases, distinctions are given that
separate the new taxa from species that can be identified using the keys presented by
Chippendale (1988), Brooker and Kleinig (1994) and Pryor et al. (1995).
Interpretation of E. kenneallyi and the extra-Australian taxa is from herbarium material
only; other taxa have been interpreted on the basis of both herbarium collections and
field observations of populations in situ.
t Deceased 1 August 1997.
504
Telopea 8(4): 2000
Section Tranversaria
Adult leaves strongly dorsiventral ('discolorous'), variously hypostomatic to partly to
± completely amphistomatic; lateral veins closely spaced, parallel, at a high angle to
the midrib (> 50°). Anthers versatile, oblong, opening by parallel slits.
The group Transversaria as defined here comprises 23 species. Most species occur in
tall, wet forests of the coast and ranges of eastern Australia, with one species in wet
forests of south-western Western Australia, one extending to New Guinea and three
endemic to islands of south-eastern Indonesia and East Timor
Species of this group are abundant around Port Jackson, and were consequently
recognised very early in the history of Australian botany. Four species were described
before 1800 by Smith (1790, 1795, 1797). Nine species were recognised by Bentham
(1867), in six different groups (six of these were species of Transversaria sens, strict.,
placed by him in three subseries of his series Normales). He also reduced £. punctata
DC. to synonymy with E. tereticornis Sm.
Maiden ( Crit. Revis. Eucalyptus 1903-1933, 6: 355) recognised 17 species, placing them
(with other taxa) in section Macrantherae subsection Tereticornes (excepting £. pellita
F. Muell., which he placed in subsection Longiores series Non-Corymbosae). He went on
to place most of these in his seed series Lepidotae-Fimbriatae. Neither Maiden nor
Bentham considered critical features of leaf morphology in their classifications.
Blakely (1934) recognised a series Transversae based on leaf venation, and made up of
all then known taxa that we now place in Transversaria. Twenty species were included,
several of which have since been reduced to synonymy. Blakely also included
£. cladocalyx F. Muell. The latter was excluded from the series by Pryor and Johnson
(1971), who otherwise included the same taxa as Blakely. Nomenclatural problems
with taxa in the Indonesian Islands were resolved with the description of E. urophylla
by Blake (1977), and further taxonomic difficulties in this group were resolved by
Pryor et al. (1995), with the recognition of two additional species (see Table 1).
Chippendale (1988) used six series names to cover the group, including E. longifolia
Link & Otto with the grey gums (the £. pimctata group; series Punctatae herein).
We now regard the E. longifolia group as sectionally distinct, and recognise five series
within the section Transversaria as here circumscribed (Table 1). Brooker (2000)
segregated the Western Australian species E. diversicolor into a separate section based
largely on positioning of valves in fruit, and coined the sectional name Latoangulatae
for tire remaining taxa included here in Transversaria.
Table 1. A classification of section Transversaria
Section Transversaria
Series Diversicolores
E. diversicolor
Series Deaneanae
E. brunnea
E. deanei
Series Salignae
Subseries Salignosae
E. grandis
E. saligna
E. botryoides
Hill and Johnson, Eucalyptus (Myrtaceae)
505
Subseries Robustosae
E. robusta
Series Resiniferae
E. urophylla
E. orophila
E. wetarensis
E. biterranea
E. pellita
E. scias
subsp. tanyula
subsp. scias
subsp. callimastha
E. notabilis
E. resinifera
subsp. hemilampra
subsp. resinifera
E. macta
Series Punctatae
Subseries Propinquosae
E. major
E. propinqua
Subseries Punctatosae
E. biturbinata
E. canaliculata
E. punctata
E. grisea
E. longirostrata
Series Resiniferae
Bark persistent or mostly so, shortly fibrous. Disc raised or level, rarely depressed.
1. Eucalyptus biterranea L.A.S. Johnson & K.D. Hill, sp. nov.
Ab E. pellita distinguitur: alabastra, fructus et folia minora, fructus saepissime angustiores.
Type: Queensland, N. slope of Mt Tozer (12°45'S, 143°12E), K. Hill 1862, P. Hind &
D. Healey, 28 July 1986 (holo NSW; iso BRI, CANB, PERTH).
Tree to 25 m tall. Bark persistent to larger branches, long fibrous-flaky, reddish-brown
underneath, smooth, grey above. Juvenile leaves disjunct, ovate acuminate, petiolate.
Adult leaves lanceolate to broad lanceolate, dorsiventral, hypostomatic except for a
few scattered stomata adjacent to the adaxial midrib, disjunct, 70-200 mm long, 20-50 mm
wide; petioles 12-35 mm long; lateral veins closely-spaced, regular, at 45°-60°to
midrib, closely and regularly reticulate between; intramarginal vein distinct, 0.5-2 mm
from margin. Inflorescence simple, axillary; umbellasters 7-flowered. Peduncles
flattened, 9-18 mm long, to 3 mm wide at apex. Pedicels terete or angular, 4-12 mm
long. Mature buds ovoid to broadly fusiform, 14-18 mm long, 7-9 mm diam.; calyptra
506
Telopea 8(4): 2000
1.5-2 times longer than hypanthium, inflated conical, shortly and broadly rostrate.
Stamens all fertile, filaments erect in bud, anthers oblong, dorsifixed, versatile,
dehiscing by parallel slits. Fruits cup shaped, 3- rarely 4-5-locular, 7-11 mm long, 7-11 mm
diam.; hypanthium smooth; calyptra scar and stemonophore broad, raised; disc
depressed; valves vertically exserted, triangular, apiculate.
Previously confused with and included in E. pellita, which we now regard as an
Australian endemic restricted to the Cairns region. £. biterranea is distinguished from
£. pellita by the smaller buds, fruits and leaves (Table 2). It is the only species in section
Transversaria occurring in New Guinea, readily distinguished from other eucalypt
species in the region by the strongly discolorous leaves with closely parallel pinnate
venation, and the fruits with prominently exserted valves. This is the taxon referred to
as species A by Pryor et al. (1995), who also discuss diagnostic features of the
Indonesian and Timorese taxa. It is distinguished from £. orophila and £. ivetarensis by
the calyptra more than 1.5 times the length of hypanthium in bud (about equal in the
other species), and from £. urophylla by the broad fruits, about as long as wide (fruits
are longer than wide in £. urophi/lla).
Table 2. A comparison of the tropical Australian red mahogany species
leaves, I x b mm
petiole mm
peduncles mm
pedicels mm
buds, I x b mm
fruits, I x b mm
E. biterranea
70-200 x 20-50
12-35
9-18
4-12
14-18x7-9
7-11x7-11
E. pellita
110-200x35-75
18-25
15-25
5-10
13-16x7-10
10-13x9-18
E. macta
70-180x12-35
15-25
5-25
3-12
14-21 x 6-8
7-10x7-10
E. resinifera
90-220 x 20-45
15-30
9-20
4-10
14-20 x 5-6
6-10x6-9
istribution: sporadic in distribution, through the Macllwraith and Iron ranges and
northwards in Australia, the southern margin of the Oriomo Plateau in the Western
Province of Papua New Guinea, and the south-east of the Merauke region in Irian Jaya
(Fig- 1)-
Ecology: usually a tree of rainforest margins.
Conservation status: not considered to be at risk.
Etymology: from the Latin bi-, two, and terra, a land or region, in reference to the
species occurrence in both far north Queensland and the island of New Guinea (Papua
New Guinea and Irian Jaya).
w!r!vcur K '7 nS (fr0m 15 examined): Australia: Queensland: Cape Flattery, Bean 469, 24 June
nrrL c r km W ° f Barrow Point, Cape York Peninsula, Cape Melville National Park, Fell
innc tlcunt" 21 JU ’ y 1993 ( BRI ' CANB, NSW); Mcllwraith Range, Hyland 7659, 22 Sep 1974
(C ,NSV\ " T R ' 14 ' Mdlwraith Range, Leo Ck road, Hyland 8411, 22 Sep 1975 (QRS, NSW); Tozers
Gap, Irvine 227, 30 June 1972 (QRS, NSW).
G . uinea: Westem Province: Approx. 6 km S of Keru on road to Mata, Gunn 838,22 Sep
tNSW); Approx. 1.2 km S of Kumbalusi to(wards) Mata, Gunn 849, 850, 22 Sep 1987 (NSW);
QM/ P Tr : 2 °rr eS North ofTokwa on road to Kiriwo, Sirisa, Gunn 865, 871, 29 Sep 1987 (NSW);
bw ot Coe (half a days walk or about 9 km), 12 km N of Kiriwo, Gunn 882, 885, 1 Oct 1987 (NSW);
new track to mining camp S of Maru, McDonald 837, 23 Oct 1988 (CANB, NSW).
Indonesia: Irian Jaya: Merauke: Djidjurug, Long BW569 (NSW).
Hill and Johnson, Eucalyptus (Myrtaceae)
507
Fig. 1. Distribution of E. biterranea, E. orophylla, E. pellita, E. wetarensis, E. urophylla.
2. Eucalyptus macta L.A.S. Johnson & K.D. Hill, sp. nov.
Ab E. resinifera distinguitur: calyptra diametro majore, brevior, plus rostrata; fructus major,
latior et relative brevior; discus descendens, valvae robustiores et prominenter exsertae.
Type: Queensland: Wild River, 2.5 km east of Herberton on Atherton road. Hill 1115,
Johnson & Blaxell, 15 Aug 1984 (holo NSW; iso BRI, CANB, DNA).
Tree to 25 m tall. Bark persistent to smallest branches, long fibrous-flaky,
reddish-brown underneath. Juvenile leaves disjunct, ovate, acuminate, petiolate.
Adult leaves lanceolate, dorsiventral, hypostomatic except for a few scattered stomata
adjacent to the adaxial midrib, disjunct, 70-180 mm long, 12-35 mm wide; petiole
15-25 mm long; lateral veins closely-spaced, regular, at 45°-60°to midrib, closely and
regularly reticulate between; intramarginal vein distinct, 0.5-2 mm from margin.
Inflorescence simple, axillary; umbellasters (7-)ll-flowered. Peduncles flattened, 5-25 mm
long, to 4 mm wide at apex. Pedicels terete or angular, 3-12 mm long. Mature buds
ovoid to fusiform, 14-21 mm long, 6-8 mm diam.; calyptra 2-3 times longer than
hypanthium, usually broadly rostrate for about V 2 length. Stamens all fertile,
filaments erect in bud, anthers oblong, dorsifixed, versatile, dehiscing by parallel slits.
Fruits cup-shaped, 3-4-locular, 7-10 mm long, 7-10 mm diam.; calyptra scar and
stemonophore broad, steeply raised, 1-1.5 mm wide; disc depressed or level and
incurved, 1-2 mm wide; valves vertically exserted, triangular, apiculate.
E. macta is distinguished from E. resinifera Sm. by the broader, shorter, more rostrate
calyptra, and the larger, broader and relatively shorter fruit with a descending disc
and heavier, more prominently exserted valves (Table 2).
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Distribution: Queensland: Sporadically distributed in highland areas of North
Queensland, the Macllwraith range, Atherton Tableland and Eungella range (Fig. 2).
Ecology: a component of wet sclerophyU forests in sheltered gullies on sandy soils
over granite, sandstone or acid volcanic rocks. Associated species include E. tereticomis,
E. eugenioides Sieber ex Sprengel, Corymbia tessellnris (F. Muell.) K.D. Hill & L.A.S.
Johnson and C. intermedia (R. Baker) K.D. Hill & L.A.S. Johnson.
The epithet is from the Latin mactus, magnified, referring chiefly to the fruit characters
as compared to those of E. resinifera.
Conservation status: not considered to be at risk.
Selected specimens (from 19 examined): Queensland: 8 miles [12.8 km] from Atherton towards
Herberton, Brooker 3393 ,28 Jan 1972 (CANB, NSW); ridge between Herberton and Atherton, Brooker
4123 ,23 Aug 1973 (CANB, NSW); 7 km along Bennetts Road, NW of Eungella, Brooker 5250 ,21 July
1996 (CANB, NSW); forestry road, west of Atherton to Herberton road, Brooker 6510, 9 Oct 1979
(CANB, NSW); west of Paluma, Croivley 158 ,11 Feb 1985 (NSW); 8 km east of Hidden Valley on
Paluma road. Hill 1154, Johnson & Blaxell, 18 Aug 1984 (NSW, BRI, CANB, PERTH); Herberton
Range, Hyland 5142 ,7 June 1971 (QRS, NSW); SFR 144, Hyland 5699 ,16 Nov 1971 (QRS, NSW); SFR
194, Hyland 5718, 2 Dec 1971 (QRS, NSW); Trnaroo Hills, Hill 1148 & Johnson, 17 Aug 1984 (NSW);
tablelands, Herberton district, Mocatta 16, Dec 1915 (NSW); Portion 42, Tarzali, Stocker 1496 ,30 July
1976 (QRS, NSW); 1.3 km SE of Wallum Trig, Herberton Range, on forestry track, Weston 1800 &
Brown, 8 July 1994 (NSW); Mt Spurgeon, White 10701, Sep 1936 (BRI, NSW).
Fig. 2. Distribution of E. macta, E. resinifera.
Hill and Johnson, Eucalyptus (Myrtaceae)
509
Series Punctatae
Bark smooth, shedding irregularly in scales or large flakes over several years, leaving
a dull surface becoming granular with age. Juvenile leaves petiolate, opposite for 4-6
nodes. Adult leaves dorsiventral, variously hypostomatic to amphistomatic.
Umbellasters 7-or-more-flowered. Disc broad, raised.
3. Eucalyptus grisea L.A.S. Johnson & K.D. Hill, sp. nov.
Inter subseriem Punctatosas distinguitur combinatione characterum sequentium: folia
juvenilia adultaque lata, calyptra moderate rostrata, pedunculus valide applanatus.
Type: Queensland: Vicinity of Pumphole Spring N of Dooloogarah Creek, NW of Mt
Moffatt homestead, Martensz 1131 & Johnston, 3 Feb 1977 (holo NSW; iso AD, BRI,
CANB, MEL).
Tree to 28 m. Bark smooth, patchy, dark grey brown, grey, orange, cream and whitish.
Seedling leaves opposite for c. 3-4 nodes. Juvenile leaves disjunct, hypostomatic,
oblong-ovate, apically rounded, to 100 mm long, 30-50 mm wide, petiolate. Adult
leaves disjunct, hypostomatic, ovate-lanceolate, acuminate, sometimes falcate,
100-180 mm long, 30-60 mm wide, petioles 15-40 mm long. Lateral veins not closely
spaced, anastomosing, at 45-60° to midrib, intramarginal vein distinct, about 1 mm
from margin. Inflorescence simple, axillary; umbellasters 7- flowered. Peduncles
strongly flattened, to 4 mm wide, 10-20 mm long. Pedicels angled, 2-5 mm long.
Mature buds oval, 6-8 mm long, 4-5 mm in diameter. Calyptra as long as hypanthium,
rounded and shortly rostrate, outer calyptra scar present as a more or less distinct lip.
Fruit cup-shaped, 8-10 mm long, 7-12 mm in diameter. Calyptra scar a shallow groove
c. 0.5 mm wide, stemonophore a narrow ridge. Disc slightly raised or flat, ultimately
incurved and enclosing valve bases, 1.5—3 mm wide. Valves broadly triangular,
apiculate, exserted and incurved at c. 45°. Seeds glossy, brown, angular, cuboid, c. 1 mm
long, chaff similar, longer and narrower.
E. grisea is distinguished in the subseries Punctatosae by the combination of the broad
juvenile and adult leaves, the beaked calyptra and the strongly flattened peduncle.
Distribution: Queensland: Consuelo Tableland, Carnarvon Range, west of Bundaberg
(Fig. 3).
Ecology: locally common in tall woodland or open forest on basalt-derived clay loam,
in association with £. melanophloia F. Muell., £. tereticornis, E. laevopinca R. Baker,
E. melliodora A. Cunn. ex Schauer and Corytnbia erythrophloia (Blakely) K.D. Hill &
L.A.S. Johnson. This species is notable in that it grows on deep heavy clay soils over
basalt, in marked contrast to other members of the Punctatosae. In some areas of this
species range, however, basalts and sandstones are closely associated, and
characteristically sandstone species such as Corymbia hendersonii K.D. Hill & L.A.S.
Johnson grow in close proximity.
Conservation status: not considered to be at risk.
The epithet is from the late Latin, griseus, 'grey', referring to the bark of the mature tree.
Selected specimens (from 14 examined): Queensland: top of The Knoll, E of Van Dyke Creek,
Buckland Tableland, Bean 799, 16 Apr 1988 (NSW); between Kookaburra Cave turnoff and
Consuelo Tableland, NW of Injune, Brooker 4869, 28 Apr 1975 (CANB, NSW); approach from west
to Consuelo Tableland, Carnarvon National Park, Brooker 4874, 28 Apr 1975 (CANB, NSW); Great
Dividing Range, c. 80 km SW of Rolleston, Peawaddy Gorge lookout, Crisp 3049, 15 June 1977
(CBG, BRI, CANB, L, NSW); 0.3 km N of 3rd crossing of Dawson River, 60 km N of Injune, Hill
4815, 1 Oct 1996 (NSW, BRI, CANB); c. 3 km N of Pumphole Spring & Dooloogarah Creek, Great
Dividing Range, Martensz 1137, 1138 & Johnston, 4 Feb 1977 (CANB, BRI, MEL, NSW); c. 6 km N
of Dooloogarah homestead and N of Dooloogarah Creek, Great Dividing Range, Martensz 1158 &
Johnston, 5 Feb 1977 (CANB, BRI, MEL, NSW).
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Section Exsertaria, the Red Gums
Cotyledons shallowly obreniform. Juvenile leaves linear to ovate, petiolate, opposite
for 3-6 nodes (more in neotenous species of series Albae). Adult leaves similifacial,
venation open, reticulate, more than one oil gland per vein island. Oil glands not
present in pith. Bark shedding regularly, or irregularly in ± large flakes, or persistent,
shortly fibrous-flaky. Calyptra clearly divided into calycine and corolline segments,
calycine calyptra shed very early or retained until near anthesis in some members of
series Bancroftianae. Filaments erect or inflexed in bud. Anthers obovate, dorsifixed,
versatile, cells distinct, dehiscing through long parallel slits. Gland relatively large,
ovate or orbicular. Filaments gently apically tapered to a fine point attached near
gland. Disc lobed in bud, inner face free from ovary; or disc absent in bud, upper wall
of ovary differentiating to form a raised, disc-like structure in fruit. Ovules usually in
6 vertical rows on placenta, rarely 4 or 8. Seeds variably shaped, variably reticulate,
reticulum sharp-edged; hilutn ventral or terminal; testa single or double.
The 'Red Gums' are a complex group comprising four series, parts of series Albae and
Brevifoliae being discussed here. Tire red gum group is difficult to diagnose, and not
unequivocally monophyletic. Possible synapomorphies for the group are the
narrowed nectary disc, the proliferation of rows of ovules on the placenta, and the
openly reticulate venation with several oil glands per vein island. Within the group,
series Tereticornes and Bancroftianae show apomorphic seed coat characters, and the
erect filaments and terminal hilum may be synapomorphies uniting the two groups.
Series Albae shows variation in seed coat character, although within a relatively
narrow range, and series Brevifoliae shows distinct apomorphic seed structure. The
Hill and Johnson, Eucalyptus (Myrtaceae)
511
very thinly peeling bark leaving a powdery salmon or orange fresh surface is uniform
in the two groups, and may be a synapomorphy, as may also the disjunct, petiolate,
rounded juvenile leaves. The thickened inner calyptra is apomorphic in series Albae,
although not clearly present in all species of the series. The thickened valves are also
apomorphic in this series.
The most likely sister group to the red gums is section Transversaria, which shows
similarities in seed coat structure, the proliferation of ovules on the placenta, and the
subterminal hilum in some groups (although the latter is possibly a result of close
packing caused by the increased number of rows of ovules).
The first Red Gum to be described was E. tereticornis Sm. (1791). Bentham (1867)
recognised two Red Gum species, which he included in Series Nomales subseries
Subexserlae (together with members of Sections Transversaria, Bisectaria , Maidenaria and
Adnataria sensu Pryor & Johnson, 1971).
Maiden (Crit. Revis. Eucalyptus 1903-1933, 6) recognised five species, placed in section
Macrantherae of his anther classification. These he then placed (with many other
species) in subsection Tereticomes series Leiophloiae and Lepidophloiae, and subsection
Longiores series Non-Corymbosae.
Blakely (1934) recognised 10 species and four varieties (three of which are here treated
as species), placing them in widely different groups (Table 3).
Table 3. Placement of tropical red gum species by Blakely (1934)
Species
Section
Subsection
Series
Subseries
E. herbertiana
Macrantherae
Tereticomes
Exsertae
Phaeoxyla
E. confluens
Macrantherae
Tereticomes
Exsertae
Erythroxyla
E. brevifolia
Macrantherae
Tereticomes
Subexsertae
Argophloiae
E. alba
Macrantherae
Tereticomes
Subexsertae
Argophloiae
E. platyphylla
Macrantherae
Tereticomes
Subexsertae
Argophloiae
E. bigalerita
Macrantherae
Tereticomes
Subexsertae
Argophloiae
E. houseana
Macrantherae
Normales
Argyrophyllae
E. apodophylla
Macrantherae
Normales
Argyrophyllae
E. mooreana
E. umbrawarrensis
Macrantherae
Micrantherae
Normales
Subulatae
Argyrophyllae
Blake (1953) recognised seven species, placing them (along with many other species)
in four different series.
Pryor and Johnson (1971) grouped the red gums in section Exsertaria, with three series.
The first was series Albae, comprising eight published species and forshadowing three
new species and reduction of three of Blakely's species to subspecies. This
arrangement included all of the then known taxa except the problematical
E. umbrawarrensis Maiden, but also included the taxon later published as E. urophylla
S.T. Blake, now known to belong to Section Transversaria. All remaining red gums were
placed in series Tereticomes, and the enigmatic E. michaeliana Blakely was placed in a
third monotypic series in this section.
Chippendale (1988) followed the series of Pryor and Johnson, with the recognition of
all of Blakely's species and the inclusion of E. umbrawarrensis (following Johnson 1972),
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recognising 15 species in series Albne. This included four recently described species
(post Pryor & Johnson), and excluded £. urophylla. Other red gum series also followed
Pryor and Johnson. Brooker and Slee (1994) departed from this arrangement,
separating the E. brevifolia E MuelL group as a separate series Brevifoliae on the basis of
autapomorphic seed coat characters. Brooker (2000) presents another approach to the
red gum group, dividing it into three sections with no further comment on possible
relationships and placing £. michaeliam in a fourth section.
We now recognise that four series are discernible (Table 4), but would still regard them
as allied and falling within section Exsertaria as diagnosed above (excluding
£. michaeliam). A total of about 48 species make up the section, with 20 species in the
two series Albae and Brevifoliae. Most species in these two series occur in areas that
have until recently been poorly collected, and confusion has surrounded species
determination and nomenclature. The Eucalyptus alba group (subseries Albosae)
presents a number of taxonomic problems in Indonesia, Timor and Papua New
Guinea, and will be discussed elsewhere.
Table 4. A classification of the red gums, section Exsertaria
Section Exsertaria
Series Albae Series Bancroftianae
Subseries Albosae
E. alba
E. tintinnans
E. platyphylla
E. bigalerita
E. houseana
E. apodophylla
E. seeana
E. disdusa
E. interstans
E. prava
E. bancroftii
E. parramattensis
subsp. decadens
subsp. parramattensis
Subseries Mooreanosae
Series Tereticornae
Subseries Tereticornosae
E. amplifolia
E. mooreana
E. pantoleuca
Subseries Herbertianosae
subsp. amplifolia
subsp. sessiliflora
E. glomericassis
E. herbertiana
E. cupularis
E. gregoriensis
E. tereticornis
E. glaucina
E. kabiana
E. blakelyi
E. chlorodada
E. terrica
E. dealbata
E. dwyeri
E. vicina
E. nandewarica
E. flindersii
Subseries Hallianosae
E. hallii
Series Brevifoliae
E. rupestris
E. kenneallyi
E. umbrawarrensis
E. confluens
E. brevifolia
E. leucophloia
subsp. leucophloia
subsp. euroa
E. ordeana
Subseries Camaldulensosae
E. camaldulensis
E. obtusa
E. rudis
Subseries Exsertosae
E. brassiana
E. lockyeri
E. exserta
E. ammophila
E. nudicaulis
E. morrisii
E. gillenii
Hill and Johnson, Eucalyptus (Myrtaceae)
513
Key to the series
1 Seeds glossy red . Series Bancroftianae
1* Seeds not glossy red
2 All or some filaments erect in bud, hilum terminal . Series Tereticornes
2* Filaments not erect in bud, hilum ventral
3 Seeds brownish-black, ragged and deeply reticulate. Series Albae
3* Seeds yellow-brown or pale brown, smooth and shallowly reticulate.
. Series Brevifoliae
Series Albae
Bark wholly smooth, shed regularly in thin flakes, trunks usually at first salmon or
orange, becoming 'powdery' white. Juvenile leaves becoming disjunct very early
(except in neotenous species), broad-lanceolate or ovate to orbicular. Outer calyptra
shed early in development. Inner calyptra ± hemispherical, ± thickened. Stamens
inflexed in bud. Nectary disc absent in bud, narrowed in fruit. Valves ± thickened.
Seeds brownish-black, ragged and deeply reticulate.
Distributed almost entirely within savanna woodland country of the monsoon tropics,
a single species occurring further south on the Queensland coast (E. hnllii Brooker near
Bundaberg). Four subseries are recognised.
Subseries Albosae
Subseries Mooreanosae
Subseries Herbertianosae
Subseries Hallianosae
Juvenile leaves more or less orbicular, disjunct.
Juvenile leaves orbicular, opposite
Juvenile leaves lanceolate to ovate, disjunct.
Juvenile leaves large, lanceolate to ovate, falcate, disjunct.
Subseries Mooreanosae
Adult leaves opposite, orbiculate.
Although apparently paralleling the neotenous condition occurring in some other
eucalypt groups, this condition is not strictly neotenous in this group. In this case, the
first few pairs of juvenile leaves are opposite, broad-lanceolate and petiolate; later
juvenile leaves remain opposite but become orbiculate (and sessile in £. mooreana), and
adult leaves retain the latter condition.
3. Eucalyptus pantoleuca L.A.S. Johnson & K.D. Hill, sp. nov.
Ab E. mooreana distinguitur: alabastra, fructus et folia majora, folia adulta petiolata et
habitus arborescens in arenosis planitierum saepe inundatarum.
Type: Western Australia: 10 km W of 'Tableland' homestead, L.A.S. Johnson 2019,
21 Aug 1967 (holo NSW).
Tree to 12 m tall, usually less than 6 m, often of twisted habit. Bark smooth throughout,
powdery, white or salmon, orange or pink, shedding in large plates or flakes. Juvenile
leaves lanceolate to ovate, dull grey green, petiolate. Adult leaves opposite to
subopposite, ovate to suborbiculate or orbiculate, not falcate, obtuse or rounded,
basally rounded or cordate, dull, glaucous, coriaceous, concolorous, 90-130 mm long,
70-100 mm wide; petioles narrowly flattened, decurrent into ridges on stems, 20-30 mm
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Telopea 8(4): 2000
long; lateral veins prominent, obtuse, widely spaced, regular, reticulum complete;
intramarginal vein obscure, looped, 2-4 mm from margin. Inflorescence simple,
axillary; umbellasters 3-flowered. Peduncles terete or angular, thick, 8-20 mm long.
Pedicels angular, 0-5 mm long. Buds ovoid or globose, glaucous, 17-24 mm long,
14-20 mm diam.; calyx calyptrate; shedding early; corolla calyptrate, free from calyx;
calyptra hemispherical, smooth, about as long as hypanthium, wider than
hypanthium; hypanthium smooth or angular. Flowers cream or yellow, all stamens
fertile, filaments indexed in bud, anthers versatile, oblong, dehiscing by parallel slits;
style base not sunken. Fruits conical, pedicellate, 3-4-locular, smooth to ribbed or
angular, 18-24 mm long, 16-20 mm diam.; calyptra scars forming a flat or slightly
raised rim to 3 mm wide; stemonophore evident as a distinct narrow groove to mm
wide; disc flat or slightly raised, less than 1 mm wide; valves exserted, raised at 30-60°.
Seeds irregular, ovoid, shallowly reticulate, dull, dark brown, 1-2 mm long; hilum
ventral. Chaff dimorphic, linear and cuboid, similar in colour to seeds. (Fig. 4).
Fig. 4. £. pantoleuca. a, adult leaves, inflorescences and buds; b, inflorescence and buds; c, transverse
section of bud; d, anther; e, fruit, (from Johnson 2019). Scale bar: a, b, c, e = 10 mm; d = 0.5 mm.
Hill and Johnson, Eucalyptus (Myrtaceae)
515
E. pantoleuca is most closely allied to E. mooreana W.V. Fitzg. ex Maiden, differing in
the larger buds, fruits and leaves, the distinctly petiolate adult leaves (sessile and
sometimes connate in £. mooreana), and the flood-plain habitat.
Distribution: Western Australia, north and east Kimberley region (Fig. 5).
Ecology: sporadically distributed on sandy soils, usually on low-lying areas on
flood-plains or seasonally wet areas.
Occasional hybrids with E. camaldulensis var. obtusa are known in areas where the two
species occur in proximity.
The epithet is from the Greek, panto, 'entirely', and leukon, 'white', referring to the
distinctly glaucous, white nature of the entire plant.
Conservation status: although locally common, populations are widely separated and
erratically distributed (3R).
Selected specimens (from 18 examined): Western Australia: Gibb River Rd, Dunlop 6038 & Done,
11 Nov 1981 (DNA, BRI, CANB, DNA, MEL, NSW, PERTH); ca 8 km E of Reed Spring Yard, W
of Tableland Homestead, George 15160,18 June 1978 (PERTH, NSW); 40 km W of kununurra on
Wyndham rd. Hill 936, Johnson & Benson, 23 July 1984 (NSW); 25 km W of Durack River crossing
on Kununurra to Gibb River rd, Hill 946, Johnson & Benson, 23 July 1984 (NSW, CANB, DNA,
PERTH); 65 miles [104 km] S of Kalumburu Mission, North Kimberleys, Lazarides 4971 & Speck,
9 Sep 1954 (CANB, NSW); 11 miles [17.6 km] E of Gibb River Station, kimberleys, Lazarides 4991
& Speck, 12 Sep 1954 (CANB, NSW); 10 miles [16 km] SE of Tableland Station, Kimberleys, Lazarides
5116, 18 Apr 1955 (CANB, NSW); 6 miles [9.6 km] W of Tableland Station, Kimberleys, Lazarides
6413, 22 July 1959 (CANB, NSW); 25 miles [40 km] NE of Karunjie Station, Perry 3091, 29 July 1952
(CANB, BRI, DNA, K, MEL, NSW, PERTH, US).
Fig. 5. Distribution of E. mooreana, E. pantoleuca.
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Telopea 8(4): 2000
Subseries Herbertianosae
Juvenile and adult leaves disjunct, petiolate. Juvenile leaves lanceolate to ovate. Disc
narrow. Valves thick.
4. Eucalyptus glomericassis L.A.S. Johnson & K.D. Hill, sp. nov.
Ab £. cupulare et E. herbertianae distinguitur: folia longa, lucide viridia, alabastra saepe
verrucosa et calyptra latissima.
Type: Northern Territory: Deaf Adder Gorge, (13°02'S 132°57'E), C. Dunlop 4344,
21 Feb 1977 (holo NSW; iso DNA).
Tree to 10 m tall. Bark smooth throughout, white to grey, brown or pink, shedding in
large plates or flakes. Adult leaves disjunct, narrow lanceolate to lanceolate, often
falcate, acuminate, basally tapered, highly glossy, green, concolorous, 120-250 mm
long, 8-35 mm wide; petioles narrowly flattened or channelled, 12-27 mm long; lateral
veins prominent, acute, widely spaced, irregular; reticulum complete; intramarginal
vein distinct, continuous, 0.5-1 mm from margin. Inflorescence simple, axillary;
umbellasters 7-flowered. Peduncles terete or weakly angular, 6-16 mm long. Pedicels
terete or angular, 1-5 mm long. Buds ovoid or globose, 7-8 mm long, 5-6 mm diam.;
calyx calyptrate; shedding shortly before anthesis; corolla calyptrate, free from calyx;
calyptra hemispherical, smooth or irregularly verrucose, about as long as hypanthium,
wider than hypanthium; hypanthium smooth or verrucose. Flowers white or cream,
all stamens fertile, filaments irregularly inflexed in bud, anthers versatile, oblong,
dehiscing by parallel slits; style base not sunken. Fruits hemispherical or conical, 4-5-
locular, smooth or verrucose, 7-9 mm long, 7-9 mm diam.; calyptra scar raised at
60-90' , 0.5-1.0 mm wide, ± stepped inside hypanthium; stemonophore raised, 0.5 mm
wide; disc raised, ultimately incurved, 1.5-2.5 mm wide; valves broadly triangular,
exserted, vertically raised, somewhat incurved at tips. Seeds irregularly ovoid to
pyramidal, shallowly reticulate, semi-glossy, dark brown, 1.5-2 mm long; hilum
ventral. Chaff dimorphic, linear and cuboid, similar in colour to seeds. (Fig. 6).
£. glomericassis differs from £. cupularis C. Gardner, £. gregoriensis N.G. Walsh & D.E.
Albrecht and £. herbertiana Maiden in the long, narrow glossy green leaves and the
often verrucose buds with a broad hypanthium and an even broader calyptra.
Distribution: Northern Territory: known only from the sandstone massifs of Kakadu
and western Arnhem Land (Fig. 7).
Ecology: restricted to skeletal soil on very rugged sandstone country, often along
watercourses but not in deep gorges. Almost always near the broken eroded edges of
the plateau rather than on the relict surface of the actual plateau.
Conservation status: not considered to be at risk.
The epithet is from Latin glomus, glomeris a ball and cassis a helmet, referring to the
shape of the calyptra.
Selected specimens (from 24 examined): Northern Territory: Mt Brockman plateau Kakadu
National Park, Boland 2129, 2130, 2131,2132, 2133, 2134 & Wardman, 17 Nov 1984 (CANB, NSW);
hilltop W of Nabarlek, Arnhem Land, Brooker 5358, 2 Oct 1976 (CANB, NSW); Magela Creek,
Dunlop 3369, 25 Feb 1973 (DNA, NSW); SW Nablek, Dunlop 4289, 20 Oct 1976 (DNA, NSW); 44 km
SE Oenpelli, Dunlop 4927, 14 June 1978 (DNA, CANB, NSW); top of Jim Jim Falls, Dunlop 5660,
30 Jan 1981 (DNA, BRI, CANB, MEL, NSW); Upper Goomadeer River, Dunlop 7232, 1 Nov 1987
(DNA, BRI, MEL, NSW); Twin Falls, about 200 m back along creek above falls. Hill 903, 16 July 1984
(NSW); Amhemland Plateau, Lazarides 7546, 6 July 1972 (CANB, BRI, DNA, K, L, NSW, US); Site
64, 14.5 km NE of Jabira East, Lazarides 9010, 26 May 1980 (CANB, NSW); 1 km E of Mt Gilruth,
Olsen 2704, 2705, 5 June 1976 (NSW).
Hill and Johnson, Eucalyptus (Myrtaceae)
517
Fig. 6. E. glomericassis. a, adult leaves; b, infructescence and fruits; c, transverse section of bud;
d, anther, (a, b from Brooker 5358; c, d from Dunlop 5660). Scale bar: a = 10 mm; b = 5 mm; c = 1 mm;
d = 0.5 mm.
Series Brevifoliae
Bark wholly smooth, shed regularly in small thin scales, trunks usually at first salmon
or orange, becoming 'powdery' white or silver. Juvenile leaves becoming disjunct very
early, ovate to orbicular, petiolate. Outer calyptra shed early in development. Inner
calyptra hemispherical to conical, not thickened. Stamens irregularly flexed in bud.
Nectary disc absent in bud, narrowed in fruit. Valves thin. Seeds rounded to elliptical,
distinctly regularly shallowly reticulate, yellow-brown.
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Telopea 8(4): 2000
Fig. 7. Distribution of E. cupularis, E. glotnericassis, E. herbertiana.
A series of six close species, distributed almost entirely within savanna woodland
country of the monsoon tropics of Western Australia and the Northern Territory
(Figs. 11 & 13). See Brooker and Kleinig (1994) for keys to species.
5. Eucalyptus kenneallyi K.D. Hill & L.A.S. Johnson, sp. nov.
Ab E. rupestri distinguitur: folia latiora breviaque; ab E. umbrawarrense distinguitur:
folia non nitentia et latiora.
Type: Western Australia: Storr Island, between Doubtful Bay and George Water,
Kimberley Coast, K. Kennealh/11083 ,17 July 1990 (holo NSW; iso PERTH).
Tree to 8 m tall. Bark smooth throughout, white to grey, brown or pink, shedding in
large plates or flakes. Intermediate leaves becoming disjunct early, lanceolate to ovate,
dull grey green, petiolate. Adult leaves disjunct, narrow lanceolate to lanceolate, not
falcate, acuminate, basally tapered, dull, green to grey-green, chartaceous,
concolorous, 50-110 mm long, 7-18 mm wide; petioles terete, 13-25 mm long; lateral
veins obscure, acute, moderately spaced, regular; reticulum complete; intramarginal
vein obscure or confluent with margin, continuous, 0-1 mm from margin.
Inflorescence simple, axillary; umbellasters 7-flowered. Peduncles terete, 3-5 mm long.
Pedicels terete, 1-2 mm long. Buds ovoid to clavate, not glaucous or pruinose, 6-7 mm
long, 2.5-3 mm diam.; calyx calyptrate; shedding early; corolla calyptrate, free from
calyx; calyptra conical, smooth, about '/ 2 a s long as hypanthium, as wide as or slightly
wider than hypanthium; hypanthium smooth. Flowers white or cream, all stamens
fertile, filaments irregularly flexed in bud, anthers versatile, oblong, dehiscing by
parallel slits; style base not sunken. Fruits cylindrical, pedicellate, 3-locular, smooth,
4-5 mm long, 3-3.5 mm diam.; calyptra scar flat, 0.2 mm wide; stemonophore
depressed, 0.2 mm wide; disc depressed, not incurved, 1-1.5 mm wide; valves
enclosed, raised at 60-80° to vertically raised. Seeds regular, ovoid to globose, finely
shallowly reticulate, semiglossy, pale brown, 0.5 mm long; hilum ventral. Chaff
dimorphic, linear and cuboid, similar in colour to seeds.
Hill and Johnson, Eucalyptus (Myrtaceae)
519
Distinguished by the thin, narrow, dull, adult leaves and the very small fruits with a
depressed disc. Most closely allied to E. mpestris Brooker & Done from the central
Kimberley region, which is readily distinguished by the broader and shorter leaves.
These two taxa form a sister group to E. umbrawarrensis, a Northern Territory endemic,
which is distinguished by the narrow and highly glossy leaves
Distribution: Western Kimberley Region, known at present only from offshore islands
(Fig. 8).
Ecology: restricted to skeletal sandy soils on hard siliceous outcrops.
Conservation status: not known (2K).
The epithet refers to Kevin Kenneally, of the Department of Conservation and Land
Management, a most active and experienced investigator of many parts of the
Kimberley region of Western Australia.
Other specimen examined: Western Australia: Koolan Island, Wannan 7, Jan 1974 (UNSW, NSW).
6. Eucalyptus leucophloia Brooker, Nuytsia 2(2): 112, fig. 7 (1976).
Type: Western Australia: near Rudall River (22°37’S, 122°12'E), AS. George 10782,
22 May 1971 (holo PERTH; iso CANB, K, NSW).
Tree to 10 m tall. Bark smooth throughout, white, powdery, shedding in small dark
scales. Intermediate leaves opposite, orbiculate, dull grey-green to slightly glaucous,
petiolate, to 70 mm long, 50 mm wide. Adult leaves disjunct, narrow lanceolate to
lanceolate, not falcate, acuminate, basally tapered, dull, green to grey-green,
chartaceous, concolorous, 50-110 mm long, 8-20 mm wide; petioles terete, 13-25 mm
long; lateral veins obscure, acute, moderately spaced, regular; reticulum complete;
intramarginal vein obscure or confluent with margin, continuous, 0-1 mm from
margin. Inflorescence simple, axillary; umbellasters 7-11-flowered. Peduncles terete,
4—10 mm long. Pedicels terete, 1-2 mm long. Buds ovoid, not glaucous or pruinose,
Fig. 8. Distribution of E. kenneallyi, E. rupestris, E. umbrawarrensis.
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7-9 mm long, 3-5 mm diam.; calyx calyptrate; shedding early; corolla calyptrate, free
from calyx; calyptra hemispherical to broadly conical, smooth, about as long as or
slightly shorter than hypanthium, as wide as hypanthium; hypanthium smooth.
Flowers white or cream, all stamens fertile, filaments irregularly flexed in bud, anthers
versatile, oblong, dehiscing by parallel slits; style base not sunken. Fruits cylindrical,
pedicellate, 3-locular, smooth, 5-8 mm long, 4-6 mm diam.; calyptra scar flat, 0.5 mm
wide; stemonophore depressed, 0.2 mm wide; disc depressed, not incurved, 1-2 mm
wide; valves enclosed, raised at 60-80° to vertically raised. Seeds flattened-ovoid,
shallowly finely reticulate, semi-glossy, yellow-brown, c. 1 mm long; hilum ventral.
Chaff dimorphic, linear and cuboid, similar in colour to seeds. (Fig. 9).
Two allopatric subspecies may be recognised on differences in fruit morphology.
1 Valves mostly enclosed... 6A. subsp. leucophloia
1* Valves prominently exserted. 6B. subsp. euroa
6A. Eucalyptus leucophloia Brooker subsp. leucophloia
Distribution: restricted to the Pilbara Region of Western Australia (Fig. 10).
Ecology: widespread and common on shallower sandy soils on sandstone, usually on
rises and sloping sites.
Hybrids have been recorded with E. trivalvis Blakely.
Conservation status: Not considered to be at risk.
Selected specimens (from 16 examined): Western Australia: Radio Hill, Paraburdoo (Paraburdoo
is c. 60 km S of Tom Price), Boomsma 657 ,10 July 1980 (AD, NSW); Dale's Gorge, Hamersley Range,
SE of Wittenoom, Briggs 3609, 15 June 1970 (NSW); Millstream, 90 miles [144 km] NW of
Wittenoom, Brooker 2079, 24 Sep 1969 (PERTH, NSW); 54 miles [86.4 km] N of Shepherds
Roadhouse towards Roy Hill, Brooker 4555, 21 Apr 1974 (CANB, NSW); Shovelanna Creek rd,
E of Newman, Brooker 8191, 4 July 1983 (CANB, NSW); At Red Gorge, between Tom Price and
Wittenoom, 18 km S of Wittenoom, in Hamersley National Park, Croat 52273A, 6 Aug 1981 (MO,
NSW); Wittenoon [Wittenoom] Gorge, above the old asbestos mine, Croat 52280A, 6 Aug 1981 (MO,
NSW); Wittenoom Gorge, Hamersley Ranges, Johnson 2124, 28 Aug 1967 (NSW); Mt Florence
[Florance] near Roebourne, McVicar 1819, 24 Aug 1922 (NSW); Tributary of Watrara Creek, Maslitt
2068, 3 Sep 1971 (PERTH, NSW).
6B. Eucalyptus leucophloia Brooker subsp. euroa L.A.S. Johnson & K.D. Hill, subsp. nov.
Ab subspecie leucophloia distinguitur: valvae manifeste exsertae.
Type: Northern Territory: 34 km S of Macarthur R crossing on Tablelands Highway, K.D.
Hill 1023, L.A.S. Johnson & D. Benson, 6 Aug 1984 (holo NSW; iso CANB, DNA, PERTH).
[E. leucophloia subsp. QQ Brooker & Kleinig 1994)]
Readily distinguished by the steeply raised and prominently exserted valves.
Distribution: across the drier monsoon tropics of the Northern Territory and
Queensland, from Top Springs east to Cloncurry, and from Daly Waters south to
Wauchope (Fig. 10).
Ecology: widespread in open savanna woodlands on shallow sandy soils on sandstone.
Some intergrades with E. brevifolia occur to the west of Top Springs in the Northern
Territory.
Conservation status: not considered to be at risk.
The epithet is from the Latin eurous, eastern, referring to its occurrence in the eastern
part of the species range.
Hill and Johnson, Eucalyptus (Myrtaceae)
521
Fig. 9. E. leucophloia subsp. euroa. a, juvenile leaves; b, adult leaves, inflorescences and buds;
c, transverse section of bud; d, anther; e, fruits; f, seed. E. leucophloia subsp. leucophloia.
g, infructescence and fruits, (a from Hill 1023; b, c, d from NSW 132433; e, f from Hill 889; g from
Croat 52273A). Scale bar: a, b = 10 mm; e, g = 2 mm; c, f = 1 mm; d = 0.5 mm.
Selected specimens (from 63 examined): Queensland: Alexander River Crossing on the
Cloncurry/Burketown rd, Carolin 8848, 24 Apr 1974 (SYD, NSW); 60.4 km from Adel's Grove,
towards Gregory Downs, 32.1 km W of Gregory River, Dalliston HC313, 6 Aug 1987 (BR1, NSW);
11.7 km N of Daly Waters turn off on Stuart Highway, Hill 889, Johnson & Benson, 13 July 1984
(NSW); 60.9 km N of Burke and Wills roadhouse on Normanton rd, Hill 1047 & Johnson, 9 Aug 1984
(NSW); 10 miles [16 km] NW of Cloncurry Township, Lazarides 4310, 11 Mar 1954 (CANB, NSW);
40 km W of Stuart Highway on rd to Top Springs, leg. ign. NSW 304159, 1 July 1978 (NSW);
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Camooweal Caves National Park, McDonald CC9, 12 Aug 1987 (BRI, NSW); c. 3 km SE of
'Wernadinga' Station, Pullen 9013, 2 May 1974 (CANB, NSW); 105.4 km W of Normanton on
Burketown rd, Puttock 12584 & St George, 11 Nov 1982 (UNSW, CANB, DNA, NSW, QRS); 64 miles
[102 km] SE of Burketown Township, Speck 4775,25 July 1954 (CANB, NSW); Spring Creek 27 km
N Mt Isa on Camooweal rd. Turner 62 & Connell , 10 Aug 1977 (CANB, BRI, DNA, MEL, NSW); Mica
Creek 15 km S Mt Isa, Turner 66 ,11 Aug 1977 (CANB, BRI, DNA, MEL, NSW); Lawn Hill, 'Cascade
Track', Williams 84101 ,15 Aug 1984 (BRI, NSW).
Northern Territory: Mt Isa turn-off from Stuart Highway, 622 miles [1001 km] S of Darwin,
Beaugleholc 10703 ,16 July 1965 (MEL, CANB, DNA, NSW); 3.3 km N of Tennant Creek, Brooker 9973,
22 June 1988 (CANB, NSW); 15 miles [24 km] W of Wollogorang on road to Calvert hills, Carolin
9224 ,12 May 1974 (SYD, NSW); Devil's Marbles, Chippendale, 28 Mar 1956 (DNA, NSW); 31.2 miles
[50 km] SE of lop Springs store, Chippendale & Johnson, 4 Oct 1957 (DNA, NSW); Benners Springs,
Forde 230 ,5 July 1956 (DNA, NSW); 5 km along road from Barkly Tableland Highway to Kilgour
Gorge, Halford 8458 ,19 May 1984 (DNA, DNA, NSW); 3.2 km S of Old Highland Plains Homestead,
Henry 247 ,26 July 1971 (DNA, NSW); 2.9 km N of Wauchope on Stuart Highway, Hill 876, Johnson
& Benson, 12 July 1984 (NSW, CANB, DNA, PERTH); 61 miles [98 km[ S Tennant Creek, Newtown,
16 Aug 1963 (DNA, NSW); 47 miles [75 km] E 3 Way Road House, Nicholls 588, 30June 1967 (DNA,
NSW); 63 miles [101 km] W of Frewena, Olsen 454, 1 July 1967 (NSW); 20 miles [32 km] SSE of
Willeroo Station, Perry 2846 ,30 May 1952 (CANB, NSW); Sunday Creek/Kalala Boundary, Sivertsen
947 ,18 Aug 1984 (DNA, CANB, DNA, MEL, NSW); Tire Pebbles, 7 km W of Stuart Highway, c. 17
km NNW of Tennant Creek, Wilson 4539 ,19 Apr 1983 (NSW).
Section Adnataria
Bark smooth or partly or fully persistent, shortly fibrous and flaky ('box'), or ± corky,
± kino-impregnated ('ironbark'). Pith glands present or absent. Cotyledons rounded
or shallowly obreniform. Seedling leaves petiolate, opposite for few, rarely many
nodes. Inflorescences sometimes simple, more often compounded with aggregations
of unit umbellasters on short, leafless lateral or apparently terminal shoots
(pseudoterminal). Compounded inflorescences usually do not continue vegetative
growth on the same axis. Calycine calyptra shed early or persisting to anthesis, ± free
from coralline calyptra; or fused to coralline calyptra. Filaments irregularly flexed or
indexed in bud, rarely erect (E. fibrosa). Anthers adnate, basifixed, opening by slits or pores.
Fig. 10. Distribution of E. leucophloia subsp. leucophloia, subsp. euroa, E. ordensis.
Hill and Johnson, Eucalyptus (Myrtaceae)
523
Section Adnatnrin is a large and widespread group, occurring through all of Australia
except Tasmania and the far south-west of Western Australia. The group is probably
monophyletic, defined by the adnate anthers and petiolate early seedling leaves.
Maximum diversity is reached in eastern Australia, with the greatest of infrasectional
variety. A total of about 125 species in about 14 series can be recognised; only series
Oligatlthae being discussed here. The nature and definition of several of the possible
series requires further study, and no key to the series is presented.
Series Oliganthae
Bark persistent, shortly fibrous and flaky ('box'), sometimes shedding on branches or
throughout in subseries Microthecosae. Seedling leaves petiolate, opposite for few
nodes (many in subseries Pruinorosae). Adult leaves similifacial, densely regularly
reticulate, lateral veins regular, at 40-60° to midrib. Pith glands absent. Inflorescences
axillary or on leafless lateral or apparently terminal shoots (pseudoterminal),
compounded with aggregations of unit 7-flowered umbellasters; or irregular in
subseries Microthecosae (units are not regularly umbelliform, but have varying degrees
of intemode development). Calycine calyptra free from corolla, shed before anthesis.
Stamens all fertile, filaments irregularly flexed in bud. Anthers adnate, globular to
oblong, opening by broad lateral pores or short slits. Fruit persistent or dehiscing
immediately on maturing. Seeds elliptical, shallowly reticulate; hilum ventral.
Defined within the section by the regular leaf venation, the free calycine calyptra
shedding before anthesis, the absence of staminodes, and the irregularly flexed
filaments with more or less globose anthers. These characters are probably all
plesiomorphic within Adnataria, but the Oliganthae also appear to share a common
biogeographic history.
Series Oliganthae was first recognised as a distinct taxonomic group in 1971 (Pryor &
Johnson 1971). The first taxa in the group to be described were E. oligantha and
E. pruinosa, named by Schauer (1843). These were followed by E. leptophleba,
E. microtheca, E. patellaris and E. spodophylla (a later synonym of E. pruinosa), named by
Mueller (1859).
Bentham (1867) recognised five species, placing them in series Porantherae (E. pruinosa
and E. oligantha), series Micrantherae (E. microtheca, mistakenly as E. brachypoda, and
E. leptophleba), and series Normales subseries Subexsertae (E. patellaris).
Species of this group were included in anther series Porantheroideae by Maiden
(Crit. Revis. Eucalyptus 1903-1933, 6: 529), together with species from many other
groups. Maiden later (Crit. Revis. Eucalyptus 1903-1933, 7:121) placed these species in
seed series Striolatae, again with taxa from very many other groups.
Blakely (1934) recognised 14 species, which he placed in section Macrantherae, series
Paniculatae subseries Oliganthae (E. argillacea), section Porantheroideae series Buxeales
subseries Subplatyphyllae, Megaphyllae, Boreales and Protrusae (most species), and series
Siderophloiae subseries ]ugatae (E. pruinosa).
Blake (1953) recognised eight species in the Northern Territory, placing seven of them
(along with E. normantonensis Maiden & Cambage) in series Buxeales, and the eighth
with the ironbarks in series Siderophloiae (E. pruinosa).
Pryor and Johnson (1971) recognised a series Oliganthae with ten species (including
E. rummeryi Maiden, now excluded, and not including E. pruinosa). This series was
included in the wider group of boxes and ironbarks, section Adnataria in subgenus
Symphyomyrtus. We later revised the circumscription of the Oliganthae to exclude
E. rummeryi and include E. pruinosa (in lists privately circulated to some eucalypt workers).
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Telopea 8(4): 2000
Chippendale (1988) adopted the series name Striolntne from Maiden, but with our
modified circumscription following Pryor and Johnson, and included 14 species.
Brooker (2000) coined the superfluous name Aquiloninres for essentially the same group.
We now recognise 32 species in this group, falling into eight subgroups or subseries
(Table 5, and see key below). Most species occur in areas which have until recently
been poorly collected, and confusion has surrounded species determination and
nomenclature. All species except some of the E. coolabah Blakely & Jacobs group are
tropical in occurrence (Hill & Johnson 1994). Members of this series may be locally
abundant, but tend not to dominate large tracts of country in the manner of, for
example, the related £. albens Benth. in New South Wales.
Table 5. A classification of the tropical boxes. Section Adnataria,
Section Adnataria
Series Oliganthae
Subseries Leptophlebosae
E. leptophleba
E. pa tel laris
Subseries Oliganthosae
E. oligantha
subsp. oligantha
subsp. modica
E. fitzgeraldii
E. koolpinensis
Subseries Tectificosae
E. tectifica
E, costuligera
Subseries Chlorophyllosae
E. chlorophylla
Subseries Argillaceosae
£ epruinata
E. microneura
E. tropica
E. leucophylla
£. limitaris
E. tephrodes
E. argillacea
E. xerothermica
series Oliganthae.
Subseries Distantosae
E. distans
E. obconica
Subseries Microthecosae
E. acroleuca
E. microtheca
E. cyanoclada
E. barklyensis
E. gymnoteles
E. helenae
E. coolabah
subsp. excerata
subsp. coolabah
subsp. arida
E. victrix
Subseries Pruinorosae
E. pruinosa
subsp. tenuata
subsp. pruinosa
Key to the subseries
1 Fruits persistent, more or less woody
2 Adult leaves disjunct, petiolate
3 Adult leaves ovate to orbiculate . Subseries Oliganthosae
3* Adult leaves narrowly to broadly lanceolate
4 Adult leaves glossy
5 Fruits large, with thick valves. Subseries Leptophlebosae
5* Fruits small, with thin valves . Subseries Chlorophyllosae
4* Adult leaves dull
Hill and Johnson, Eucalyptus (Myrtaceae)
525
6 Leaves more or less coriaceous, not deciduous Subseries Argillaceosae
6* Leaves chartaceous, mostly deciduous in dry season .
. Subseries Distantosae
2* Adult leaves opposite, sessile . Subseries Pruinorosae
1* Fruits dehiscing immediately upon maturity, more or less chartaceous
7 Inflorescence units regularly umbelliform . Subseries Tectificosae
7* Inflorescence units not regularly umbelliform . Subseries Microthecosae
Subseries Oliganthosae
Bark fully persistent. Adult leaves ovate to orbiculate, petiolate. Fruit persistent. Seeds
dark grey-brown or charcoal to black.
A subseries of three species occurring in the north of the Northern Territory and the
Kimberley region of Western Australia.
7. Eucalyptus oligantha Schauer, in Walpers, Repert. Bot. Syst. 2: 926 (1843).
Type: Northern Territory: Copeland Island, Mountnorris Bay, A. Cunningham 250, 12
Apr 1818 (lecto (here designated) BM; isolecto CANB, K, NSW). Cited as: A. Cunn.
Herb. no. 250/1818.' It is unclear as to where the primary set of specimens studied by
Schauer may have been, and the BM specimen is hence here designated the lectotype
since the BM was the primary repository and source for the Cunningham collections.
= E. hillii Maiden, /. & Proc. Roy. Soc. New South Wales 53: 63 (1919).
Type: Northern Territory: Bathurst Island, G.F. Hill 468, 6 Nov 1916 (holo NSW).
Included in E. oligantha by Blake (1953).
= E. hillii Maiden var. alleniana Blakely & Jacobs in Blakely, Key Eucalypts : 239 (1934).
Type: Northern Territory: near Stapleton, M.R. Jacobs 58, 12 July 1935 (holo NSW 10054).
Included in E. oligantha by Blake (1953).
Tree to 15 m tall. Bark persistent throughout, pale grey, shortly fibrous-flaky. Juvenile
leaves disjunct early, petiolate, ovate to orbiculate. Adult leaves disjunct, similifacial,
broadly ovate to orbiculate, semiglossy to glossy dark green, acute or obtuse, 50-150 mm
long, 40-130 mm wide; petioles 25-60 mm long. Lateral veins ± closely-spaced,
regular, at 40-60° to midrib; reticulum even, dense; oil glands small, dense, often
difficult to see; intramarginal vein continuous, distinct, 0.5-1 mm from margin.
Inflorescences compounded, pseudoterminal or lateral; unit umbellasters
3-7-flowered. Peduncles terete, 4-14 mm long. Pedicels terete, 2-11 mm long. Mature
buds ovoid to clavate, apiculate, smooth, 6-9 mm long, 3-5 mm diam.
Calyptra hemispherical to conical, sometimes apiculate, _-l times as long as
hypanthium. Fruit cylindrical to cup-shaped, smooth, 3-4-locular, 5-11 mm long, 5-9 mm
diam.; calyptra scar and stemonophore flat or somewhat depressed, c. 0.5 mm wide;
disc vertically depressed, 1-2 mm wide; Valves broadly triangular, obtuse, basally
enclosed, vertically raised with tips at rim level to distinctly exserted.
Two subspecies may be distinguished on regional differences in leaf, bud and fruit size.
8A. subsp. oligantha
... 8B. subsp. modica
1 Fruit large (6-11 x 6-9 mm)
1* Fruit small (5-6 x 5-6 mm).
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Telopea 8(4): 2000
8A. Eucalyptus oligantha subsp. oligantha
Adult leaves 60-150 mm long, 50-130 mm wide, petioles 25-60 mm long. Peduncles
4— 11 mm long. Pedicels 2-11 mm long. Buds 7-9 mm long, 4-5 mm diam. Fruits 6-11 mm
long, 6-9 mm diam.
Distribution: north of Pine Creek in the Northern Territory, extending to Melville
Island (Fig. 11).
Ecology: a fairly localised species, usually occurring in open savanna woodlands on
flat sites on alluvial sandy soils near creeks.
Hybrids are recorded with E. tectifica F. Muell. and E. distans Brooker, D. Boland
& D. Kleinig.
Conservation status: widespread and locally abundant, not considered to be at risk.
Selected specimens (from 24 examined): Northern Territory: just W of Adelaide R, and Margaret
R, Blrtke 16991 ,13 Sep 1946 (BRI, NSW); 1 km W of Bark Hut hotel on the Arnhem 1 Iighway, Boland
& Wardman 2110, 14 Nov 1984 (CANB, NSW); flats near Primpininga Ck N of Paru [Melville
Island], Brooker 3221, 29 June 1971 (CANB, NSW); Copeland Island, NW of Murgenella, Dunlop
6936, 21 Oct 1987 (DNA, BRI, CANB, MEL, NSW); c. 20 km from Pine Ck on Jabiru rd. Hill 900,
Johnson & Benson, 15 July 1984 (NSW, CANB, DNA, PERTH); Melville Island, Maclear Ck, S of
Pickataramoor, Hill 908 ,17 July 1984 (NSW); 10 km N of Bridge Ck on New Stuart Highway, Hill
3341, Johnson & Stanberg, 17 Nov 1988 (NSW); Bathurst Island, Jacobs 41 ,1 July 1933 (NSW); 20 miles
[33 km[ NNE of Tipperary Homestead, Lazarides 6659 ,19 July 1961 (CANB, BRI, NSW); Coomalie
Ck, Stuart Highway, Waterhouse & Wannan 11379 ,15 Jan 1981 (UNSW, NSW).
8B. Eucalyptus oligantha subsp. modica L.A.S. Johnson & K.D. Hill, subsp. nov.
Ab subspecie oligantha foliis, alabastris et fructibus totis minoribus delicatioribusque
distinguitur.
Type: Western Australia: 1 km E of Durack R. crossing on Gibb R rd, K.D. Hill 943,
L.A.S. Johnson & D. Benson, 23 July 1984 (holo NSW; iso CANB, DNA, PERTH).
Adult leaves 50-100 mm long, 40-90 mm wide, petioles 25-45 mm long. Peduncles
5- 14 mm long. Pedicels 4-7 mm long. Buds 6-8 mm long, 3-4 mm diam. Fruits 5-6 mm
long, 5-6 mm diam.
Fig. 11. Distribution of E.fitzgeraldii, E. koolpinensis, E. oligantha subsp. oligantha, subsp. modica.
Hill and Johnson, Eucalyptus (Myrtaceae)
527
Distinguished from the type subspecies by the smaller and more delicate leaves, buds
and fruits.
Distribution: north and east of Gibb River station in Western Australia, not extending
east of the Durack Range (Fig. 11).
Ecology: restricted and highly localised, usually on alluvial sandy soils near creeks.
Conservation status: although of rather restricted distribution and not abundant
within its range, this species occurs in a habitat that is not significantly threatened (3R).
The epithet is from the Latin modicus , moderate-sized, from the smaller sizes of all
parts of this subspecies in contrast to the type subspecies.
Selected specimens (from 6 examined): Western Australia: 13 km W of Berkeley R, Brooker 7778,
2 Nov 1982 (CANB, NSW); 7.2 km W of King George R crossing on track to Carson R., Hill 953,
Johnson & Benson, 25 July 1984 (NSW); 'Gibb River' Station near Homestead, Johnson 2090 ,26 Aug
1967 (NSW); 50 miles 182 km] NNW of Gibb R. Station, Speck 4977 ,10 Sep 1954 (CANB, NSW).
Subseries Tectificosae
Bark fully persistent. Adult leaves lanceolate. Fruit chartaceous, dehiscing
immediately on maturing. Seeds grey-brown to dark grey-brown.
A subseries of two species occurring in grassy savanna country in north-western
Queensland and the north of the Northern Territory and Western Australia.
Key to species
1 Fruits smooth, 4—6 mm long, 4—5 mm diam. E. tectifica
1* Fruits ribbed, 6-8 mm long, 5-7 mm diam. E. costuligera
9. Eucalyptus costuligera L.A.S. Johnson & K.D. Hill, sp. nov.
Ab E. tectifica distinguitur: folia alabastraque glauca, alabastra fructusque longiores et
costati. Habitat sola sicca, saxosa aliquantum elevata.
Type: Western Australia: 60 km E of Gibb River to Kalumburu road on Wyndham road
(16°20'S, 126°55'E), M.I.H. Brooker 7780, 3 Nov 1982 (holo NSW; iso CANB, PERTH).
Tree to 10 m tall. Bark persistent throughout, pale grey, shortly fibrous-flaky (box).
Adult leaves disjunct, similifacial, lanceolate, obtuse, 70-150 mm long, 17-30 mm
wide; petioles to 22 mm long; lateral veins ± closely-spaced, regular, at 40-60° to
midrib; reticulum even, dense; oil glands small, dense; intramarginal vein continuous,
distinct, <0.5 mm from margin or at margin. Inflorescences compounded,
pseudoterminal or axillary; unit umbellasters 1-7-flowered, somewhat irregular.
Peduncles terete, 1-15 mm long. Pedicels terete or ± ribbed, 2-6 mm long. Mature buds
clavate, apiculate, regularly shallowly ribbed; c. 7 mm long, c. 4 mm diam.; calyptra
broadly hemispherical, apiculate to rostrate, c. 1/2 as long as hypanthium. Fruit
cylindrical to cup-shaped or pyriform, apically constricted, ± chartaceous, often
regularly shallowly ribbed, 3-4-locular, 6-8 mm long, 5-7 mm diam., deciduous at
maturity; calyptra scar and stemonophore flat or somewhat depressed, <0.5 mm wide;
disc vertically depressed, 1-2 mm wide; valves broadly triangular, obtuse, wholly
enclosed, vertically raised, ± chartaceous. (Fig. 12).
Eucalyptus costuligera differs from E. tectifica F. Muell. in the more coriaceous, glaucous
leaves and buds, the longer, ribbed buds and fruit, and the differing habitat, on
elevated, dry, shallow, stony soils.
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Telopea 8(4): 2000
Distribution: Western Australia: central north-east Kimberley region (Fig. 13).
Ecology: locally abundant but apparently restricted on dry, elevated skeletal soils over
sedimentary rocks, with Corymbia grandifalia (R. Br. ex Benth.) K.D. Hill & L.A.S.
Johnson, Eucalyptus obconica Brooker & Kleinig and £. jensenii Maiden.
Conservation status: widespread and locally abundant, not considered to be at risk.
The epithet is from the Latin costa, a vein or rib, with the diminutive suffix -ula, -ulum,
together with the suffix -ger, bearing or carrying, referring to the distinctively finely
ribbed fruits.
Specimens examined: Western Australia: 11 km W of Durack R crossing on Gibb
R. rd, Hill 944, Johnson & Benson, 23 July 1984 (NSW); 12.7 km W of Durack R on Gibb R. Rd, Hill
3374, Johnson & Stanberg, 19 Nov 1988 (NSW, CANB, PERTH); 82 miles [c. 133 km] SW of Wyndham
Township, Perry 3087, 29 Aug 1952 (CANB, NSW).
Fig. 12. E. costuligera. a, adult leaves, inflorescences and buds; b, transverse section of bud;
c, anther; d, fruits, (from Brooker 7780). Scale bar: a, d = 10 mm; b = 1 mm; c = 0.5 mm.
Hill and Johnson, Eucalyptus (Myrtaceae)
529
LEGEND
♦ E. costuligera
A E. tectifica
130 ° 132 ° 134 ° 136 ° 138 ° 140 ° 142 °
I I I I I I I-
10 °
t
Fig. 13. Distribution of E. costuligera, E. tectifica.
Subseries Argillaceosae
Bark fully persistent. Adult leaves dull, narrow-lanceolate to narrow-ovate. Fruit
small to large, persistent. Seeds dark grey-brown or charcoal to black.
A subseries generally occurring in the drier parts of the monsoon savanna region,
extending onto drier sites in some wetter areas. The nine species are closely related,
and occur in a geographic replacement pattern with some overlap, where ecological
segregation is evident.
Key to species (after Brooker & Kleinig 1994)
1 Twigs, leaves and buds glaucous
2 Qld species
3 Fruits cup-shaped.
3* Fruits obconical.
2* NT and WA species
4 Fruits cup-shaped .
4* Fruits obconical .
1* Twigs, leaves and buds not glaucous
5 Qld species
6 Fruits cup-shaped .
6* Fruits obconical.
5* NT and WA species
. E. tropica
E. microneura
. E. tephrodes
.. E. argillacea
.. E. epruinata
E. leucophylla
530
Telopea 8 ( 4 ): 2000
7 Kimberley and NT; leaves thin (0.28-0.38 mm when dry); peduncles and pedicels
long (6-23 and 2-9 mm). E. limitaris
7* Pilbara; leaves thick (0.39-0.51 mm when dry); peduncles and pedicels short (2-9
and 1-3 mm) . E. xerothermica
10. Eucalyptus epruinata L.A.S. Johnson & K.D. Hill, sp. nov.
Ab E. microneum et aliis speciebus affinis distinguitur: pruina absens, petioli pedunculi
pedicellique graciles, fructus breves parvique valvis valde exsertis.
Type: Queensland: 7.6 km E of Croydon on Georgetown road (18°09'S, 142°12'E),
K. Hill 1056 & L. Johnson, 10 Aug 1984 (holo NSW; iso BRI, CANB, K).
Tree, sometimes mallee, to 6 m tall. Bark fully persistent, grey, coarsely shortly
fibrous-flaky (box). Adult leaves disjunct, similifacial, lanceolate to ovate, obtuse,
50-110 mm long, 10-35 mm wide; petioles to 22 nun long, slender; lateral veins
± closely spaced, regular, at 40-60° to midrib; reticulum even, dense; oil glands small
to large, moderately spaced; intramarginal vein continuous, distinct, 0.5-2.0 mm from
margin. Inflorescences compounded, pseudoterminal or axillary; unit umbellasters
3-/-flowered. Peduncles slender, terete, 5-15 mm long. Pedicels slender, terete, 2-6 mm
long. Fruit cylindrical to cup-shaped, 4-locular, 4-5 mm long, 4-5 mm diam.; calyptra
scar not continuous with stemonophore, flat, <0.1 mm wide; stemonophore depressed,
0.2-0.4 mm wide, with a flat, persistent staminal ring; disc flat, <0.5 mm wide, situated
below the level of the staminal ring; valves broadly triangular, obtuse or acute, fully
exserted, raised 45-90°, ± chartaceous. Seeds semi-glossy to dull, brownish-black,
rounded, elliptical, regularly shallowly reticulate; hilum ventral. Chaff orange-brown.
£. epruinata is distinguished by the absence of glaucousness, slender petiole, peduncle
and pedicel, and the small, short fruit with strongly exserted valves.
Distribution: Queensland: South western Cape York Peninsula, Croydon to
Kowanyama (Fig. 14).
Ecology: locally abundant in open woodland on flat sandy clay-loam, with E. leptophleba
F. Muell., Corymbia polycarpa (F. Muell.) K.D. Hill & L.A.S. Johnson, Melaleuca viridiflora
Sol. ex Gaertner and Petalostigma pubescens Domin.
Conservation status: not considered to be at risk.
The epithet is from the Latin pruinatus, pruinose, with the Latin prefix ex-, e-, without
or lacking, referring to the lack of pruinosity in contrast to its distinctly pruinose sister
species E. microneura Maiden & Blakely.
Selected specimens (from 6 examined): Queensland: 9.4 km W of Croydon on highway. Hill 1055
& Johnson, 10 Aug 1984 (NSW); Croydon Base Camp, Speck 4753, 21 July 1954 (CANB, NSW).
11. Eucalyptus limitaris L.A.S. Johnson & K.D. Hill, sp. nov.
Inter subseriem Argillaceosas distinguitur: folia relative tenuia non pruinosa;
pedunculae pedicellique longi; fructus mediocres, conici vel cylindrici.
Type: Western Australia: 5 km E of Mary R crossing on Fitzroy Crossing to Halls Creek
road (18°42'S, 126°54’E), K. Hill 987, L. Johnson & D. Benson, 2 Aug 1984 (holo NSW; iso
DNA, CANB, K, PERTH).
[E. sp. UU in part, Brooker & Kleinig 1994)]
Tree or mallee to 8 m tall. Bark fully persistent, deeply fissured, grey-brown, shortly
fibrous-flaky (box). Adult leaves disjunct, similifacial, dull green, narrow-lanceolate to
broad-lanceolate, 90—250 mm long, 14—35 mm wide, 0.28—0.38 mm thick; petioles
Hill and Johnson, Eucalyptus (Myrtaceae)
531
12-26 mm long; lateral veins closely-spaced, regular, at 40-60° to midrib; reticulum
even, dense; oil glands small, obscure; intramarginal vein continuous, distinct, <1 mm
from or at margin. Inflorescences compounded, pseudoterminal or lateral; unit
umbellasters 3-7-flowered. Peduncles terete, (3—)6—23 mm long. Pedicels terete or
angular, thick, 2-9 mm long. Fruit conical to cup-shaped or cylindrical, 4-5-locular,
7—11 mm long, 6-8 mm diam.; calyptra scar and stemonophore raised or flat, 0.5-0.7 mm
wide, often with a persistent staminal ring; disc flat to steeply depressed, 1-2 mm
wide; valves broadly triangular, obtuse, basally enclosed, apically rim-level or
exserted, raised at 45—90°. Seeds semiglossy to dull, brownish-black or black, rounded,
elliptical, regularly shallowly reticulate; hilum ventral. Chaff mid-brown. (Fig. 15).
E. limitaris is distinguished by the relatively thin, dull green leaves with no glaucous
bloom, the long pedicels and peduncles, and the large conical or cylindrical fruit.
Distribution: southern Kimberley region, from west of Fitzroy Crossing to Halls
Creek, and east to west of Top Springs (Fig. 16).
Ecology: locally common but sporadic, on shallow red loam over various rock types,
in open savanna shrubland, often near dry creek lines.
Pure populations occur in the west of the range around Fitzroy crossing, and again in
the east towards Top Springs. Extensive breakdown occurs however with E. tepluodes
around Halls Creek. E. limitaris and E. tephrodes display a similar extensive breakdown
pattern to that seen with the E. tropica Cambage-E. leucophylla Domin, E. xerothermica-
E. tephrodes and E. epruinata-E. microneura pairs. E. limitaris and E. tephrodes however
show extensive pure stands of the parent species, and a degree of ecological separation
Fig. 14. Distribution of E. epruinata, E. leucophylla, E. microneura, E. tropica.
532
Telopea 8 ( 4 ): 2000
P/1 VI p
Fig. 15. £. limitaris. a, adult leaves, infructescences and fruits, b, fruits; c, seed, (from Hill 987).
Scale bar: a, b = 10 mm, c = 1 mm.
in the breakdown zone. Here, E. tephrodes or plants with a greater obvious influence of
that taxon tend to favour higher ground.
Conservation status: not considered to be at risk.
The epithet is from the Latin limes, limitis, a border or boundary, in reference to the
distribution across the Western Australia-Northern Territory border.
Selected specimens (from 19 examined): Northern Territory: Montejinni turnoff on Buchanan
Highway, Hill 1007, Johnson & Benson, 4 Aug 1984 (NSW, CANB, DNA, PERTH); 53 miles [85 km]
SE of Mistake Ck Homestead, Perry 3193, 30 Aug 1952 (CANB, NSW); Mistake Ck, Robinson 88,
26 Aug 1970 (NT, NSW). Western Australia: 35 km N of highway towards Windjana Gorge, Brooker
10127, 20 Oct 1988 (CANB, NSW); 16.5 km E of Halls Ck on Nicholson rd. Hill 999, Johnson &
Benson, 3 Aug 1984 (NSW); 25.7 km E of Tunnel Ck (just E of Cajeput Ck), Hill 3443, Johnson &
Stanberg, 23 Nov 1988 (NSW, CANB, PERTH); 7.7 km from Echidna Gorge to 3 Ways, Bungle
Bungle NP, Hill 3501, Johnson & Stanberg, 28 Nov 1988 (NSW); 13.9 km N of RB creek on Duncan
Highway, Hill 3521 & Stanberg, 30 Nov 1988 (NSW, CANB, PERTH); 27 km from Duncan Highway
towards Nelson Springs, Hill 3522 & Stanberg, 30 Nov 1988 (NSW).
Hill and Johnson, Eucalyptus (Myrtaceae)
533
12. Eucalyptus tephrodes L.A.S. Johnson & K.D. Hill, sp. nov.
Inter subseriem Argillaceosas distinguitur: folia magna, crassiuscula, glauca; fructus
majusculi plus minusve aequidimensionales in pedicellis et pedunculis longis
crassisque.
Type: Western Australia: 40.1 km from highway towards Bedford Downs, K.D. Hill
3463, L.A.S. Johnson & L. Sternberg, 25 Nov 1988 (holo NSW; iso CANB, PERTH).
[E. sp. UU in part, Brooker & Kleinig 1994)|
Tree or sometimes mallee to 10 m tall. Bark fully persistent, grey to grey-white, shortly
fibrous-flaky (box). Juvenile leaves disjunct, glaucous, ovate-lanceolate to orbicular,
petiolate, to 60 mm wide, 90 mm long. Adult leaves disjunct, similifacial, glaucous,
lanceolate to broad-lanceolate, acute or rounded, 70-170 mm long, 12-45 mm wide,
0.31-0.44 mm thick; petioles 12-23 mm long; lateral veins closely-spaced, regular, at
40-60° to midrib; reticulum even, dense; oil glands small, obscure; intramarginal vein
continuous, distinct, <1 mm from or at margin. Inflorescences compounded,
pseudoterminal or axillary; unit umbellasters 3-7-flowered. Peduncles terete, 4—12 mm
long. Pedicles terete or ± flattened, 2-6 mm long. Mature buds clavate, 8-9 mm long,
4-5 mm diam.; calyptra hemispherical, shortly apiculate, c. V 2 as long as hypanthium.
Fruit cup-shaped to globular-truncate, rarely cylindrical or pyriform, 4-6-locular, 6-10
mm long, 6-11 mm diam.; calyptra scar and stemonophore flat, c. 0.5 mm wide; disc
vertically depressed,l-2 mm wide; valves broadly triangular, obtuse, deeply enclosed,
tips rarely exserted, raised at 45° or more. Seeds semi-glossy to dull, brownish-black,
rounded, elliptical, regularly shallowly reticulate; hilum ventral. Chaff orange-brown.
(Fig. 17).
E. tephrodes is distinguished by the large, moderately thick glaucous leaves and the
large more or less equidimensional fruit with long, thick pedicels and peduncles.
Leaves are thicker in the north and west of the range in the Kimberley region, and also
in the Pilbara region (0.36-0.44 mm), becoming thinner (0.31-0.37 mm) around Halls
Creek. It is more frequently a medium-size single-trunked woodland tree than the
related and partly sympatric £. limitaris. Both taxa however may form true mallees
with extensive subterranean lignotubers.
Distribution: Western Australia, south-east Kimberley region. Halls Creek and north,
with a disjunct occurrence in the eastern Pilbara region (Fig. 16).
Ecology: locally common but sporadic in open savanna woodland with Corymbia
flavescens K.D. Hill & L.A.S. Johnson, E. camaldulensis subsp. obtusa, E. brevifolia and
E. limitaris, on more or less loamy soils on various sites from dry creek-beds to low
stony hills. Quite extensive woodlands of this species occur around Bedford Downs
and west of there. In the Pilbara region, this species is less common and more widely
scattered, and occurs in red loam along drainage lines and dry watercourses in open
flat country, associated with E. trivalvis, Corymbia hamersleyana (D.J. Carr & S.G.M.
Carr) K.D. Hill & L.A.S. Johnson, E. leucophloia subsp. leucophloia, E. xerothermica and C.
Candida K.D. Hill & L.A.S. Johnson. Intergradation with E. xerothermica is discussed
under the latter. In areas where the two occur together, E. tephrodes favours flatter sites
on deeper soils, with E. xerothermica on shallower soils on higher sites. This is in
contrast to the ecological segregation between E. limitaris and E. tephrodes, where
E. tephrodes favours higher sites. The somewhat different habitat of E. tephrodes in the
Pilbara region compared with that in the Kimberley region suggests that some genetic
differences may be present, but no clear morphological distinction can be made
between the two occurrences.
Conservation status: widespread and locally abundant, not considered to be at risk.
534
Telopea 8 ( 4 ): 2000
The epithet is from the Greek tephros, ash-grey, with the Greek termination -odes, -oides,
resembling, from the grey-glaucous foliage.
Selected specimens (from 21 examined): Western Australia: c. 40 km N of Halls Ck, Beadle 179, 22
Sep 1972 (NSW); between Turkey Creek and Halls Creek, Breaker 7743, 26 Oct 1982 (CANB, NSW,
PERTH); c. 4 km NW of Mt Meharry on access track to Trig Point, Hill 483, Johnson, Blaxell, Brooker
& Edgecombe, 1 Nov 1983 (NSW, CANB, PERTH); 118.9 km W of Newman on track to Tom Price
via Mt Mehan-v, (22°56'S, 118°53'E), Hill 485, Johnson, Blaxell, Brooker, Edgecombe, 1 Nov 1983
(NSW, CANB, K, PERTH); 54 km SW of Halls Ck on highway, Hill 989, 990, Johnson & Benson,
2 Aug 1984 (NSW); 7.6 km from Bedford Downs towards Lansdowne, Hill 3470, Johnson & Stanberg,
25 Nov 1988 (NSW); Teronis Gorge at Chamberlain River crossing, Hitt 3485, Johnson & Stanberg,
26 Nov 1988 (NSW, CANB, PERTH); 6-8 km from New Halls Ck on rd to Halls Ck, Johnson 2065,
25 Aug 1967 (NSW); 0.8 km NW of Bedford Downs Homestead, Johnson 2068, 25 Aug 1967 (NSW);
14 miles [23 km] S of Rockhole Station, Lazarides 3190, 23 Aug 1952 (CANB, BRI, NSW, NT, PERTH);
16 miles [25.5 km] W of Halls Ck township. Perry 3179, 17 Aug 1952 (CANB, BRI, K, NSW, DNA,
PERTH, US).
13. Eucalyptus xerothermica L.A.S. Johnson & K.D. Hill, sp. nov.
Ab E. limitari distinguitur: folia crassiuscula; pedicellis et pedunculis brevioribus;
fructus proportione latiores.
Type: Western Australia: 'Palm Pool', c. 5 km W (downstream) of Millstream Tavern
(old homestead) on Fortescue River, K.D. Hill 430, L.A.S. Johnson, D.F. Blaxell, M.I.H.
Brooker & W. Edgecombe, 30 Oct 1983 (holo NSW; iso CANB, PERTH).
[E. sp. VV Brooker & Kleinig 1994)]
Tree or mallee to 6 m tall. Bark fully persistent, yellowish-grey to grey-brown, shortly
fibrous-flakey (box). Adult leaves disjunct, similifacial, dull green, narrow-lanceolate
to broad-lanceolate, rounded to acute, 110-230 mm long, 10-35 mm wide, 0.39-0.51 mm
thick; petioles thick, 9-21 mm long; lateral veins closely-spaced, regular, at 40-60° to
Hill and Johnson, Eucalyptus (Myrtaceae)
535
Fig. 17. E. tephrodes. a, adult leaves, inflorescences and buds; b, adult leaves, infructescences and
fruits; c, transverse section of bud; d, anther; e, fruits, (a, c, d from Brooker 7743; b, e from Johnson
2065). Scale bar: a, b, e = 10 mm; c = 1 mm; d = 0.5 mm.
536
Telopea 8 ( 4 ): 2000
midrib; reticulum even, dense; oil glands small, obscure; intramarginal vein
continuous, distinct, 0.5 mm from margin. Inflorescences simple or compounded,
pseudoterminal or axillary; unit umbellasters 3-7-flowered. Peduncles thick, terete,
2-9 mm long. Pedicels thick, terete or angular, 1-3 mm long. Fruit cylindrical to
cup-shaped, tapering into pedicel, 4-5-locular, 7-9 mm long, 7-10 mm diam.; calyptra
scar and stemonophore flat, 0.7-1.0 mm wide; disc flat to vertically depressed, 1-2 mm
wide; valves broadly triangular, obtuse or acute, basally enclosed or rim level, apically
rim level or exserted, raised at 45-90°. Seeds semi-glossy to dull, brownish black,
somewhat angular to rounded, elliptical, regularly shallowly reticulate; hilum ventral.
Chaff brown.
E. xerothennica is nearest to E. limitaris, differing in the thicker leaves, generally shorter
pedicels and peduncles, and proportionally broader fruits.
Distribution: Western Australia: Pilbara region, North West Cape and nearby islands
(Fig. 16).
Ecology: widely scattered but local, on red, shallow loam, usually calcareous and often
with calcrete, over various rock types. A component of low open savanna, associated
with £. camaldulensis var. obtusa, E. victrix L.A.S. Johnson & K.D. Hill, Corymbia
hamersleyana and C. Candida.
E. xerothermica and E. tephrodes show a pattern of intergradation similar to that of
E. limitaris and £. tephrodes, discussed above under E. limitaris. In this case, however,
E. tephrodes tends to occur on deeper soils on lower sites or on flatter country
somewhat to the east of the range of £. xerothermica. Very little pure E. tephrodes with
no E. xerothermica influence is known, however.
Conservation status: widespread and locally abundant, not considered to be at risk.
Tire epithet is from the Greek xeros, dry, and thermos, hot, referring to the hot and dry
climate of the Pilbara region.
Selected specimens (from 13 examined): Western Australia: 13 miles [21 kmj F, of Mt Bruce in
Dales Gorge, Brooker 2175 ,29 Sep 1969 (PERTH, NSW); 58.8 km S of Exmouth on main rd. Hill 404,
Johnson, Blaxell, Brooker & Edgecombe, 28 Oct 1983 (NSW); 3.5 km W of Exmouth along Charles Knife
rd, 30 km S of Exmouth, Hill 411, Johnson, Blaxell, Brooker & Edgecombe, 28 Oct 1983 (NSW, CANB,
PERTH); 35 km NW of Highway on Onslow rd. Hill 427, Johnson, Blaxell, Brooker & Edgecombe,
29 Oct 1983 (NSW); top of Mt Meharry, Hill 479, Johnson, Blaxell, Brooker & Edgecombe, 1 Nov 1983 (NSW).
Subseries Pruinorosae
Bark fully persistent, 'box'. Adult leaves opposite, ± sessile, ovate, cordate, strongly
glaucous. Fruit persistent. Seeds dark grey-brown or charcoal to black.
14. Eucalyptus pruinosa Schauer, in Walpers, Rep. Bot. Syst. 2: 926 (1843).
Type: Queensland: Sweers, Bentinck & Allen's Is., R. Brown & F. Bauer 33, Nov 1802
(holo W; iso BM, E, K). Cited as: 'In Nova Hollandia fructiferam legit Ferd. Bauer!'
Tree or mallee to 6 m tall. Bark fully persistent, grey to grey-brown, shortly
fibrous-flaky. Juvenile leaves opposite, at first petiolate, later sessile, ovate to elliptical.
Adult leaves opposite, sessile, often amplexicaul, similifacial, dull grey-green or
glaucous, ovate to elliptical, rounded to acute, 25-150 mm long, 15-90 mm wide.
Inflorescences compounded, pseudoterminal; unit umbellasters 3-7-flowered.
Peduncles terete, 4-25 mm long. Pedicels terete, 2-22 mm long. Buds narrowly ovoid
to fusiform, often glaucous, 6-12 mm long, 3-6 mm diam.; calyptra conical and acute
to hemispherical and rostrate, 1/2-1 times as long as hypanthium. Fruit cylindrical to
cup-shaped, tapering into pedicel, 3-5-locular, 5-12 mm long, 4-10 mm diam.;
calyptra scar and stemonophore flat, 0.7-1.0 mm wide; disc flat to vertically depressed,
Hill and Johnson, Eucalyptus (Myrtaceae)
537
1-2 mm wide; valves broadly triangular, obtuse or acute, basally enclosed or rim level,
apically rim level or exserted, raised at 45-90°. Seeds semi-glossy to dull, brownish
black, somewhat angular to rounded, elliptical, regularly shallowly reticulate; hilum
ventral. Chaff brown.
A widespread and locally dominant tree across a major part of the monsoon tropics,
but not including Cape York Peninsula. Two geographically separate subspecies are
recognised.
14A. Eucalyptus pruinosa Schauer subsp. pruinosa
Adult leaves 50-150 mm long, 30-90 mm wide. Peduncles 6-25 mm long, 1-2 mm
diam. Pedicels 2-8 rarely to 22 mm long. Buds 7-12 mm long, 4-6 mm diam.; calyptra
hemispherical, rostrate, 1/2-1 times as long as hypanthium. Fruits 7-12 mm long,
6-10 mm diam.
Hybrids are recorded with E. tropica, E. limitaris and £. microtheca.
Distribution: the gulf district of north-western Queensland, extending west through
the Northern Territory at corresponding latitudes, and into the south-eastern
Kimberley region of Western Australia (Fig. 18).
Ecology: a locally dominant species in low open savanna woodlands or shrublands,
often the only tree species in the community, frequently with Triodia understorey.
Substrate varies from sand to sandy loam, often with laterite and usually on more or
less flat but well-drained sites.
Conservation status: widespread and locally abundant, not considered to be at risk.
Selected specimens (from 39 examined): Northern Territory: 132.3 km SE of Fitzroy Crossing
towards Halls Creek, Brooker 70784, 24 Apr 1991 (CANB, DNA, NSW, PERTH); 7 miles [11.2 km]
NW Alexandria HS, Chippendale NT 7947, 22 Mar 1956 (DNA, NSW); 8.6 miles [12.2 km] W of
Soudan HS, Chippendale NT 3829 & Johnson, 2 Oct 1957 (DNA, NSW); 79.1 miles [126.5 km] NE
Tanami, Chippendale NT 5722, 13 Apr 1959 (DNA, NSW); 10 km S of Ti Tree, Fox s.n., June 1986
(NSW301476); Robinson River station, Gittins 7307, June 1967 (NSW); 110 km SW of Hooker Creek
toward Tanami, Gittins 2352, Aug 1971 (NSW); 3 km along track to Wollogorang from 12 Mile
Waterhole, Halford 847720, 29 Nov 1984 (DNA, NSW); 8.4 km south of Banka Banka on Stuart
Highway, Hill 887, Johnson & Benson, 13 July 1984 (NSW, CANB); 6 miles [9.6 km] NW of the
Fig. 18. Distribution of E. pruinosa subsp. pruinosa, subsp. tenuata.
538
Telopea 8(4): 2000
Granites Township, Laiarides 6256, 22 Apr 1957 (CANB, NSW); 15 miles [24 km] NW of Limbunya
station. Perry 2312, 30 June 1949 (CANB, NSW); between Renner's [Renner] Springs and Powells
Creek, White 7, 30 May 1922 (NSW).
Queensland: Camooweal, Blake 8849, 1 May 1935 (BRI, CANB, NSW, NT); Dalgonally to Fort
Constantine, Blake 12646, 20 Aug 1936 (BRI, NSW); 7 miles [11.2 km] N of Normanton on the
Karumba road. Carotin 8722, 18 Apr 1974 (NSW); Sandy Creek, 2 miles [3.2 km] from Nardoo
towards Gregory Downs, Carolin 8884, 26 Apr 1974 (NSW); Normanton to Karumba road between
Normanton and Maggieville, Clarkson 2696 b, 6 Nov 1979 (BRI, CANB, NSW); Riversleigh Holding
[c. NE of Camooweal], Cittins 800, June 1963 (NSW); 27 miles [43.2 km] NW of Cloncurry
township, Lazarides 4307, 11 Mar 1954 (CANB, NSW); 37 miles [59.2 km] of Croydon township.
Speck 4762, 21 July 1954 (CANB, NSW); about 25 miles [40 km] W [i.e. NW] of Croydon, Stocker
741, 3 June 1971 (QRS, NSW).
15B. Eucalyptus pruinosa Schauer subsp. tenuata L.A.S. Johnson & K.D. Hill, subsp. nov.
Ab subspecie pruinosa distinguitur: alabastra minora, pedicelli graciles delicatique et
fructus parvi.
Type: Northern Territory:15 1cm E of Timber Creek roadhouse on highway, K.D. Hill
919, L.A.S. Johnson & D. Benson, 21 July 1984 (holo NSW; iso CANB, DNA, PERTH).
Adult leaves 25-110 mm long, 15-60 mm wide. Peduncles 5-18 mm long, less than 1 mm
diam. Pedicels 4-8 mm long. Buds 6-8 mm long, 3—4 mm diam.; calyptra conical,
acute, about as long as hypanthium. Fruits 5-8 mm long, 4-6 mm diam.
Distinguished from the type subspecies by the small buds, the slender, delicate
pedicels and the small fruits.
Distribution: northern, wetter parts of the Northern Territory and corresponding
northern parts of the Kimberley region of Western Australia (Fig. 18).
Ecology: habitat is generally similar to that of the type subspecies, although often on
more sloping country and on a wider range of soils, some quite heavy. Habitats of this
subspecies tend to be locally drier sites with low open woodland or shrubland, in a
region of taller and more closed vegetation.
Conservation status: widespread and locally abundant, not considered to be at risk.
Tire epithet is from the Latin tenuis, slender, with the Latin adjectival termination -atits,
in reference to the slender pedicels and peduncles.
Selected specimens (from 25 examined): Northern Territory: Roper Valley station, Bateman 11,
11 June 1950 DNA, NSW); 15 miles [24 km] W of Katherine towards Wyndham, Brooker 3142,
17 June 1971 (CANB, NSW); 12m [19.2 km] west of Katherine Gorge, Brooker 3154, 17 June 1971
(CANB, NSW); Saddle Creek, 71 miles [113.6 km] by road from Timber Creek Police Stn towards
Kununurra, Brooker 4205, 29 Oct 1973 (CANB, NSW); 1 km W of Elsey Creek on Roper Road, Brooker
6213, 5 Apr 1979 (CANB, NSW); 17 miles [27.2 km] NE of Mainoru, Cole 81 & Provan, (CANB,
NSW); 14 miles [22.4 km] NNE Mainoru, Maconochie 1430, 14 June 1972 (DNA, NSW); Maude Creek
area. Speck 1621, 3 Sep 1961 (CANB, NSW); 13 miles [20.8 km] S of Delamere station. Perry 2850,
31 May 1952 (CANB, NSW); Canopy Rock, Mountain Valley Stn, Swinbourne 682, 25 Mar 1963
(NSW); 50 km S Mataranka, Stuart Highway, Wightman 1504, 2 July 1984 (DNA, CANB, K, NSW).
Western Australia: Radigans Bore [near] Bow River, Sainty & Lillyman s.n., 9 June 1982 (NSW);
Telegraph Creek about 160 km NNE of Halls Creek, Briggs 3677, 19 June 1970 (NSW); 10 km SW
Mt Barnett St [station], Kimberleys, Guymer 606, 6 Sep 1976 (BRI, NSW).
Intergrades between the subspecies
Selected specimens (from 12 examined): Northern Territory: 95.7 km SE of Fitzroy Crossing,
Ngumban Cliff area, Brooker 10782, 24 Apr 1991 (CANB, NSW, PERTH); 20 miles [32 km] SW
Borroloola, Hill 566, 2 Sep 1911 (NSW); 6.0 km W of Montejinni turnoff on Buchanan Highway, Hill
1009, Johnson & Benson, 4 Aug 1984 (NSW).
Hill and Johnson, Eucalyptus (Myrtaceae)
539
Western Australia: Rosewood station. Beard 5664, 29 June 1968 (PERTH, NSW); Mt Nyulasy, 115 km
S of Kununurra - Wyndham rd on Halls Creek rd, Hill 933, Johnson & Benson, 22 July 1984 (NSW);
93 km SE of Fitzroy Crossing on highway. Hill 984 Johnson & Benson, 2 Aug 1984 (NSW, CANB,
DNA, PERTH); opposite Kununurra Airstrip, Palzer 713, 16 Sep 1966 (CANB, NSW); near Rexona
Bore, 26 miles [41.6 km] SSE of Cherrabun station, Lazarides 6480, 14 Aug 1959 (CANB, NSW).
Acknowledgments
Our colleague in the overall project, Don Blaxell, has contributed helpful discussions
and has been involved in the recognition of several of the taxa. Thanks are also due to
David Mackay and Lesley Elkan, who prepared the illustrations, and Leonie Stanberg,
who has provided extremely valuable technical and editorial assistance throughout
the project, and prepared the maps.
References
Bentham, G. (1867) Flora Australians, vol. 3 (L. Reeve: London).
Blake, S.T, (1953) Botanical contributions of the Northern Australia Regional Survey. Studies of
northern Australian species of Eucalyptus. Austral. J. Bot. 1:185-352.
Blake, S.T.(1977) Four new species of Eucalyptus. Austrobaileya 1(1): 7
Blakely, W.F. (1934) A Key to the eucalypts (The Worker Press: Sydney).
Briggs, J.D. & Leigh, J.H. (1996) Rare or Threatened Australian Plants, revised edn. (CSIRO:
Collingwood, Vic.)
Brooker, M.J.H. (2000) Anew classification of the Genus Eucalyptus L.Her. (Myrtaceae). Austral. Syst.
Bot. 13: 79-148.
Brooker, M.I.H. & Kleinig, D.A. (1994) Field Guide to Eucalypts, vol. 3. Northern Australia (Inkata
Press: Chatswood).
Brooker, M.I.H. & Slee, A.V. (1994) Eucalyptus series Brevifoliae (Myrtaceae), a new series of northern
Australian eucalypts. Nuytsia 9: 307-314.
Chippendale, G.M. (1988) Eucalyptus. In George, A.S. (ed.), Flora of Australia, vol. 19 (AGPS:
Canberra).
Hill, K.D. & Johnson, L.A.S. (1994) Systematic studies in the eucalypts. 6. A revision of the
coolibahs. Eucalyptus subgenus Symphyomyrtus section Adnataria series Oliganthae subseries
Microthecosae (Myrtaceae). Telopea 5: 693-702.
Hill, K.D. & Johnson, L.A.S. (1995) Systematic studies in the eucalypts. 7. A revision of the
bloodwoods, genus Corymbia (Myrtaceae). Telopea 6:185-504.
Johnson, L.A.S. (1972) Evolution and classification in Eucalyptus. Proc. Linn. Soc. Neiv South Wales
97:12-29.
Maiden, J.H. (1903-1933) A critical revision of the genus Eucalyptus. 8 vols. (Govt. Printer: Sydney).
Mueller, F. (1859) Monograph of the Eucalypti of tropical Australia. /. Linn. Soc., Bot. 3: 94.
Pryor, L.D., Williams, E.R. & Gunn, B.V. (1995) A morphometric analysis of Eucalyptus urophylla and
related taxa with descriptions of two new species. Austral. Syst. Bot. 8(1): 57-70.
Pryor, L.D. & Johnson, L.A.S. (1971) A classification of the eucalypts (ANU Press: Canberra).
Schauer, J.C. (1843) Ordo LXXXV11. Myrtaceae R.Br. In Walpers, G.G. (ed.) Repertorium botanices
systemalicae ... 2, suppl. 1:920-933.
Smith, J.E. (1790) Botany. In J. White, Journal of a Voyage to New South Wales (T. Wilson: London).
Smith, J.E. (1795) Zoology and Botany ofNeiv Holland (J. Sowerby: London).
Smith, J.E. (1797) Botanical characters of some plants of the order of Myrti. Trans. Linn. Soc. London
3: 25-288.
Manuscript received 14 December 1999
Manuscript accepted 15 May 2000
.
s.
'
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TELOPEA 8(4): 407-539 • July 2000
ISSN 0312-9764
Contents
A new species of Minima (Asteraceae: Astereae) from
New South Wales P.S. Short and J.R. Hosking 407
A revision of Davidsonia (Cunoniaceae)
Gwen J. Harden and John B. Williams 413
Seedling characteristics in the Casuarinaceae
Robin Hwang and John G. Conran 429
The taxonomy of Xanthosia hnegelii and closely related species
(Apiaceae: Hydrocotyloideae) J-M. Hart 441
Actinotus periculosus (Apiaceae): a new perennial species from
eastern Australia M.J. Henwood 455
The lichen genera Phaeographis and Phaeographina (Graphidaceae)
in Australia 1: Species based on Australian type specimens
Alan W. Archer 461
Hopkinsiaceae and Lyginiaceae, two new families of Poales
in Western Australia, with revisions of Hopkinsia and Lxjginia
Barbara G. Briggs and L.A.S. Johnson 477
Systematic studies in the eucalypts. 10. New tropical and subtropical
eucalypts from Australia and New Guinea ( Eucalyptus , Myrtaceae)
K.D. Hill and L.A.S. Johnson 503
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